HARVARD UNIVERSITY
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Comparative Zoology
Volume 141, no. I, 1998
ISSN 0040-74
Tijdschrift
voor
Entomologie
A journal of systematic and evolutionary
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Netherlands Journal of Entomology
Published by the Netherlands Entomological Society
Tijdschrift voor Entomologie
A journal of systematic and evolutionary entomology since 1858
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Ontwerpers B.V., Aad Derwort, 's-Gravenhage
-*Y
Daniel J. BICKEL
Australian Museum, Sydney, NSW, Australia
AUSTRALIAN, MELANESIAN AND MICRONESIAN
ACROPSILUS MIK (DIPTERA: DOLICHOPODIDAE)
Bickel, D. J, 1998. Australian, Melanesian, and Micronesian Acropsilus Mik (Diptera:
Dolichopodidae).- Tijdschrift voor Entomologie 141: 1-17, figs. 1-21. [issn 0040-7496]. Pub-
lished 30 November 1998.
The Australian, Melanesian, and Micronesian species of the genus Acropsilus Mik (Diptera:
Dolichopodidae) are revised, and comprise eleven species: Acropsilus protractus Robinson
(Solomon Islands, New Guinea, trop. Australia), and ten newly described species: A. albitibia
(trop. Australia, New Guinea, Solomon Islands, Vanuatu), A. boharti (Irian Jaya, Solomon Is-
lands,) A. colmani ( Papua New Guinea), .A. kuranda (Queensland,), A. malaita (Solomon Is-
lands,), A., maprik (Papua New Guinea), A. nigricornis ( Solomon Islands, New Guinea, Queens-
land), A., putosa (Papua New Guinea), A. toma (Papua New Guinea), and A. udot (Micronesia:
Truk). Some species are widely distributed in the Australasian tropics.
Acropsilus is known only from the Old World, and is particularly rich and often abundant in
moist tropical habitats. The entire genus is reviewed and redefined, and its phylogenetic posi-
tion is discussed. Acropsilus is close to the Oriental genus Griphomyia and both genera are ten-
tatively referred to the Peloropeodinae, a subfamily which requires redefinition.
D.J. Bickel, Entomology Section, Australian Museum, 6 College Street, Sydney, nsw 2000
Australia. E-mail: danb@amsg.austmus.oz.au
Key words. - Diptera, Dolichopodidae, Acropsilus, Australia, Melanesia, Micronesia.
The genus Acropsilus Mik comprises small-sized
dolichopodids. Although this genus initially appears
to be rather nondescript, males of many species are
easily recognised by their elongate ivory-white cerei,
which project out from a pedunculate hypopygium.
Closer examination reveals a number of uniquely de-
rived features which define the genus, and it has not
been readily placed in any of the traditional
dolichopodid subfamilies.
Previously six species had been described from wi-
dely separated locales in Europe, Tadzhikistan, West
Africa, the Seychelles, Sumatra and the Solomon Is-
lands. However, mass trapping methods, particularly
malaise and water traps, reveal the genus to be abun-
dant and rich in the Indo-Malayan and Australasian
wet tropics. This paper considers the fauna of Aus-
tralia, Melanesia and Micronesia, and treats eleven
species, ten of which are newly described. Two
species, Acropsilus protractus and A. albitibia are wide-
spread throughout Melanesia and northern Australia
(figs. 20, 21).
Materials and acknowledgements
This study is based on material from major world
collections. Institutional abbreviations fot material
cited follows, and I thank the respective curators and
support stafTfor the loan of specimens.
AMS Australian Museum, Sydney - M. Moulds;
ANic Australian National Insect Collection, csiro,
Canberra - P. Cranston;
BPBM Bernice P. Bishop Museum, Honolulu - N.
Evenhuis;
cas California Academy of Sciences, San Francisco
- P. Arnaud;
CMNH Carnegie Museum of Natural History, Pitts-
burgh - C. Young;
CNC Biosystematics Research Institute, Agricul-
ture Canada -J. Cumming;
NMWC National Museum of Wales, Cardiff. - J.
Deeming;
USNM National Museum of Natural History,
Smithsonian Institution, Washington, D. C
- F. C. Thompson;
ZMUC Zoological Museum, University of Copen-
hagen - V. Michelsen.
S. Bullock drew the genitalic figures. C. E. Dyte
provided notes and specimens. P. Grootaert provided
valuable comments on an early draft of the manu-
script. This research was supported by Australian Bi-
ological Resources Study (a.b.r.s.) grants 85/0921
and 87/5905.
Tijdschrift voor Entomologie, volume i4i, 1998
Methods
Drawings of genitalia were made with a camera lu-
cida attached to a compound microscope. The left
lateral view of the hypopygium or male genital cap-
sule is illustrated for most species. In describing the
hypopygium, 'dorsal' and 'ventral' refer to morpho-
logical position prior to genitalic rotation and flexion.
Thus, in figures showing a lateral view of the hypopy-
gium, the top of the page is morphologically ventral,
while the bottom is dorsal. Morphological terminolo-
gy follows McAlpine (1981) and Cumming et al.
(1995). Common features are listed in the introduc-
tory descriptions and are not repeated in species de-
scriptions unless needing clarification. Measurements
are in millimeters and were made on representative
dry specimens. Body length is measured from the
base of the antennae to the tip of the seventh abdom-
inal segment. Wing length is the perpendicular dis-
tance to the apex from an imaginary extension of the
humeral crossvein; wing width is measured from the
junction of R, with the costa to the opposite side of
the wing, perpendicular to the wing's long axis. The
CuAx ratio is the length of the m-cu crossvein/distal
section CuA. The position of features on elongate
structures such as leg segments are given as a fraction
of the total length, starting from the base. The relative
lengths of the podomeres should be regarded as repre-
sentative ratios and not measurements. The ratios for
each leg are given in the following formula and punc-
tuation: trochanter + femur; tibia; tarsomere 1/2/3/
4/5.
The following abbreviations and terms are used:
Mssc - Male secondary sexual character(s), the non-
genitalic characters found only on the male body; I,
II, III: pro- , meso-, metathoracic legs; C, coxa; T,
tibia; F, femur; ac, acrostichal setae; ad, anterodorsal;
av, anteroventral; dc, dorsocentral setae; dv, dorso-
ventral; hm, postpronotal setae; np, notopleural setae;
pa, postalar setae; pd, posterodorsal; pm, presuturai
supra-alar setae; pv, posteroventral; sa, postsutural
supra-alar setae; sr, presuturai intra-alar setae; t, tar-
sus; t15, tarsomeres 1 to 5.
Systematic part
Genus Acropsilus Mik
AcropsilusMik (1878: 6). Type species: Chrysotils nigerLoew
1869: 298, monotypy.
Description
Male. - Body length ranges from 1.0-2.3 mm, but
most species about 1.3-1.6.
Head. - Dorsal postcranium flat to slightly con-
cave; male eyes distinctly separated by face and
clypeus; eye facets enlarged anteriorly and ventrally;
clypeus usually with four setae: pair midcentral short-
er setae, and pair of stronger distal setae (as in fig. 1 5);
strong verticals and strong diverging ocellars present;
postverticals represented by 4-5 short setae, continua-
tion of postorbitals; first flagellomere variably covered
in microtrichia; arista apical, and varies from arising
on entire outer curvature of first flagellomere (fig. 15)
to arising within distinct indentation on the outer
curvature (fig. 4); arista with tiny pubescence, and
longer than head height; proboscis (fig. 18) with 6
geminate pseudotracheae; epipharyngeal armature
with 2 prongs, and dorsal projection with is weakly
sclerotised, and adjacent to which are three thorn-like
setae.
Thorax. - Usually rather short, and ratio of length/
width about 1.3 (fig. 17); proepisternum with some
short setae near posterior margin, and strong seta just
dorsad of CI; posterior slope of mesonotum slightly
flattened but not depressed; ac absent; 5 dc present,
comprising 4 stronger posterior dc which decrease in
size anteriorly, and with additional very weak dc an-
teriormost, on anterior slope of thorax; 1 pa, 2 sa, an-
terior much shorter than posterior, 2 sr, 2 npl, 1 hm,
1 pm present; posterior mesonotum with distinct
transverse fold just anteriad of scutellum; median
scutellar seta strong, lateral present as weak tiny hair;
postscutellum with distinct median longitudinal
mound.
Legs. - Mostly yellow; CI and CII with short pale
anterior hairs; CII and CIII each with short lateral
seta; leg I usually without distinctive setation; FII
without distinct true subapical anterior seta, although
sometimes some subapical setae many appear slightly
stronger; FII with subapical pv seta; Til with strong
ad-pd pair at Va, strong ad at %, and with strong ven-
tral apical, and with coronet of short setae; Fill with-
out subapical anterior seta, but with weak subapical
pv seta; Till with short ad-pd pair at Va, and often
with dorsal setae on distal half; male Hit, sometimes
somewhat swollen with ventral row of hairs (mssc).
Wing (fig. 16). - Membrane hyaline; R2+3 ends in
anterior margin at Vö; R4t5, just anterior to apex; R„+5
and M subparallel; M without flexion or "bosse
alaire"; anal vein absent; anal angle weak; CuAx ratio
near 0.3.
Abdomen. - Usually dark brown; tergal setae not
strongly developed; male tergum 1 wide (fig. 17), and
distinctly wider than thorax, especially noticeable in
dried specimens; hypopygium pedunculate (fig. 1);
segment 7 prolonged with distinct elongated tergum
and sternum separated by membrane; sternum 8 with
distinctive inverted Y-shaped carina ; hypopygial fo-
ramen left basolateral; hypandrium and aedeagus
both short and emerging at angle near base of epan-
drium; hypandrium broad and apically upcurved,
Bickel: Australasian Acropsilus (Dolichopodidae)
and basally fused with epandrium; aedeagus rather
short and slightly curved, and often with notched
phallus; ventral margin of epandrium with 2 epandri-
al setae; epandrial lobe raised from ventral margin
and externally overlapping digitiform surstylar arms;
subepandrial sclerite well-developed, curved and rest-
ing between cerei, and often with group of strongly
pedunculate distal setae; cercus usually white-ivory
coloured and subtriangular, and bearing pale setae.
Female. - Similar to male except lacking mssc and
as noted: face and clypeus more widely separated, and
facets of uniform size; clypeus always with 4 setae; in
species where male first flagellomere deeply incised,
female less incised; Hit, unmodified; female tergum 1
not noticeably wide; oviscapt (figs 3, 4) divided into
two hemitergites, each with crest of four spine-like se-
tae, and bearing elongate digitiform process and
stalked group of pedunculate setae.
General remarks
Acropsilus is a distinctive dolichopodid genus known
only from the Old World. Based on the species treat-
ed below and descriptions, the genus has the following
set of diagnostic characters: clypeus of females and
most males with four projecting setae: pair midcentral
shorter setae, and pair of stronger distal setae; arista
apical; posterior slope of mesonotum slightly flat-
tened but not depressed; ac absent; 5 dc present; pos-
terior mesonotum with distinct transverse fold just
anteriad of scutellum; R4+5 and M subparallel; male
tergum 1 wide, and distinctly wider than thorax; hy-
popygium pedunculate: segment 7 elongate; sternum
8 with distinctive inverted Y-shaped carina; hypan-
drium basally fused to epandrium; epandrial lobe
raised from ventral margin and externally overlapping
digitiform surstylar arms; subepandrial sclerite well-
developed, curved and resting between cerei, and of-
ten with pedunculate distal setae; cercus usually
white-ivory coloured and prominent; oviscapt divid-
ed into two hemitergites, each with a crest of 4 spine-
like setae, and bearing elongate digitiform process
and stalked group of pedunculate setae.
Acropsilus is particularly rich in the moist Afrotrop-
ical, Oriental and tropical Australasian regions, where
many species undoubtedly await description. Howev-
er, species are also known from western Europe and
even at elevations of 1 300 m in Tadzhikistan. Speci-
mens are frequently taken in malaise and water (pan)
traps, especially in tropical lowlands. In the Ivory
Coast, Coutourier (1978) noted that A. eburneensis
was abundant throughout the forest-savannah mosaic
during the rainy season, but confined to gallery fo-
rests during the dry season. Similarly, in northern
Australia, Acropsilus species are abundant and wide-
spread during and after monsoonal rains, but are con-
fined to permanent water during the drier periods.
Males of many species have bright ivory-white cer-
ei, which may function in sexual signalling. Here it
should be noted that Acropsilus albitibia has ivory-
white tibiae I and II (mssc), which possibly augment
the function of the cerei. However, P. Grootaert
(pers. comm.) notes that many undescribed species
from Thailand have short cerei and greatly enlarged
hypopygia.
In the Australasian tropics, Acropsilus protractus
and A. albitibia are particularly widespread through-
out Melanesia and northern Australia (figs. 20, 21).
Three species occur in Australia, where the genus is
confined to the monsoonal tropics and the Queens-
land coast south to the Tropic of Capricorn. Acrop-
silus is known as far east as Vanuatu, and considering
its presence in the isolated Micronesian Truk Group,
it probably is readily dispersed.
Checklist
Acropsilus includes the following species:
albitibia sp. n. Australia (Queensland, Northern
Territory, Western Australia), Indonesia (Irian
Jaya), Papua New Guinea, Solomon Islands,
Vanuatu
boharti sp. n. Indonesia (Irian Jaya), Solomon
Islands
colmani sp. n. Papua New Guinea
eburneensis Couturier, 1978: 222. Ivory Coast
errabundus Lamb, 1922: 403. Seychelles
igori Negrobov, 1 984: 1111. Tadzhikistan
kuranda sp. n. Australia (Queensland^
malaita sp. n. Solomon Islands
maprik sp. n. Papua New Guinea
minutus Hollis, 1964: 276. Indonesia (Sumatra).
niger Loew, 1869: 298 (Cbrysotus). Hungary,
western Europe, northern Africa, St. Helena I
nigricornis sp. n. Solomon Islands, Australia
(Queensland), Indonesia (Irian Jaya )
protractus Robinson, 1963: 830. Solomon Islands,
Australia (Queensland, Northern Territory,
Western Australia), Indonesia (Irian Jaya), Papua
New Guinea
putosa sp. n. Papua New Guinea
toma sp. n. Papua New Guinea
udotsp. n. Micronesia (Truk)
The protractus group of species {Acropsilus protrac-
tus, A. boharti, A. colmani, A. kuranda, A. malaita,
and A. minutus) all have an incised, pubescent first
flagellomere. I have seen additional species from the
Philippines (bpbm), Sabah and Peninsular Malaysia
(ams) and Taiwan (cmnh) which have antenna and
hypopygia similar to A. protractus, and the group is
rich in the Oriental-Australasian tropics. Possibly the
West African A. eburneensis aiso belongs in this group.
Tijdschrift voor Entomologie, volume 141, 1998
Acropsilus errabundus from the Seychelles is distinc-
tive for being longer (body length 2.3-2.4, whereas
most Acropsilus are 1.3-1.6), and the male has elon-
gate legs and thorax, giving it a rather gracile and al-
most medeterine appearance. As well, the male It, has
a ventral row of short setae, not found in other Acrop-
silus, and it has eyes which join across the face so that
the characteristic clypeal setae absent (they are present
on the female, however). Finally, Fill of at least the
male has an isolated anterior seta at %, which I regard
as a Mssc, not a true anterior preapical seta. I have ex-
amined a male paratype from the Seychelles, Mahé
(bmnh) and males from Silhouette, Corgate Ridge,
500 m (ams).
I have not seen specimens of the Palaearctic Acrop-
silus igori. However, Negrobov's (1984) description
does not note the presence of setae on the clypeus,
nor the presence of a Y-shaped carina on sternum 8.
As well, the hypopygial figure does not indicate the
prominent cerei which are so distinctive on con-
geners. Possibly this species belongs elsewhere.
Morphological notes
Acropsilus is a well-defined genus, and most species
conform to the general description given above. Mor-
phological characters which relate to the taxonomie
placement of Acropsilus are considered below.
I. Autapomorphies
1. Male sternum 8 has a distinctive inverted Y-
shaped carina (e.g. fig. 1). This carina is present on all
the species I have examined, and is an autapomorphy
for the genus. I know of no other dolichopodid genus
with a similar structure.
2. The epandrial lobe in most Acropsilus species lies
subparallel with and externally overlaps the surstyli
(fig. 1).
3. The male tergum 1 is distinctly wider than suc-
ceeding segments, and wider than the thorax (fig. 17),
on all species.
II. Other Character States
4. The subepandrial sclerite between the cerei is
prominent, and often has a group of strongly pedun-
culate distal setae (e.g. fig. 1). Various other dolicho-
podid subfamilies (e.g. Dolichopodinae) also have
well-developed subepandrial sclerites.
5. The clypeus of females and usually males has four
projecting setae: pair midcentral shorter setae, and
pair of stronger distal setae (as in fig. 15). Strikingly,
the New World Atlantic coastal Nanomyina barbata
(Aldrich) has similarly positioned setae (see fig. 5, Ro-
binson & Vockeroth 1981). Nanomyina also has an
apical arista, but has 2 rows of ac and a rather short
hypopygium (I have not seen specimens). This config-
uration of clypeal setae therefore is probably homo-
plasious. As well, some other dolichopodid genera also
have clypeal setae but in different configurations, for
example, as a female secondary sexual character (fssc),
in species close to Syntormon flexible Becker, and in
some undescribed Australian Sympycnus species.
6. Posterior mesonotum has a distinct transverse
fold just anteriad of scutellum. This structure was not-
ed by Ulrich (1980), and also occurs in Diaphorinae.
Systematic position
Acropsilus has been assigned variously to the Sym-
pyeninae, Peloropeodinae, or left incertae sedis. Of all
established subfamilies, Acropsilus superficially ap-
pears to be close to the Medeterinae, with which it
shares postabdominal similarities (an elongate seg-
ment 7 forming an external peduncle for the hypopy-
gium), apical arista, hypandrium basally fused to
epandrium, and reduced leg setation. However, Acro-
psilus lacks the two strong apomorphic features which
define the Medeterinae: postcranium dorsally con-
cave and posterior mesoscutum strongly flattened.
Therefore it should be kept out ofthat subfamily.
In a recent paper, Grootaert & Meuffels (1997) de-
scribed the Oriental genus Griphomyia, and noted its
similarity to Acropsilus. The two genera share a num-
ber of characters: body length mostly less than 2 mm;
dorsal postcranium flat; male eyes distinctly separated
by face and clypeus; strong verticals and strong di-
verging ocellars present; thorax rather short and wide;
posterior slope of mesonotum slightly flattened but
not depressed; 5 dc present; femora II and/ or III with
rather weakly developed subapical anterior seta; legs
weakly bristled; similar venation; hypopygium pe-
dunculate; abdominal segment 7 prolonged, and sep-
arated into distinct tergum and sternum; hypandrium
and aedeagus both short and emerging at an angle
near base of epandrium; subepandrial sclerite (= 'dor-
sal appendage' in Grootaert & Meuffels 1997) well-
developed; female oviscapt divided into two hemiter-
gites, each with crest of spine-like setae and with
elongate digitiform process subtended by peduncu-
late setae.
In summary, there are strong similarities among
G riphomyia and Acropsilus, in overall habitus (see fig.
1 in Grootaert & Meuffels 1997), female oviscapt,
and especially the male postabdominal structure
(compare figures in this paper with those in Grootaert
& Meuffels 1997). The obvious differences between
the two genera (aristal position, presence/absence of
acrostichals, clypeal setae, and anal vein) are minor
and not of subfamilial significance. The two genera
might be regarded as sister taxa and certainly belong
in the same subfamily. Grootaert & Meuffels place
both in the Peloropeodinae. However, I am not en-
tirely sure of the definition of the Peloropeodinae,
and Peloropeodes itself has a strikingly different male
Bickel: Australasian Acropsilus (Dolicbopodidae)
postabdomen to the two genera considered here.
Only provisionally would I place Acropsilus and Gri-
phomyia in the Peloropeodinae, pending further in-
vestigation ofthat subfamily.
Key to male Acropsilus from Australia,
Melanesia and Micronesia
The following small-sized dolichopodid taxa are
sympatric with Acropsilus in the Australasian region,
and can be distinguished as follows:
A. Cryptophleps species also have an apical arista
and lack ac setae, but all have a broken vein M, lack
the m-cu cross vein, and have an encapsulated hy-
popygium.
B. Sympycninae of similar size have strong anterior
preapical setae on FII and Fill, ac setae usually pre-
sent, often a distal ad setal comb on TI, a dorsal arista
and an encapsulated hypopygium.
C. Medeterinae also have a pedunculate hypopy-
gium and apical arista, but have a distinctly concave
dorsal postcranium, a strongly flattened posterior
mesonotum, and lack the wide male tergum 1 .
This key to Acropsilus is difficult and most species,
apart from a few with obvious mssc, require a genital-
ic preparation for accurate identification.
1. Cercus davate, distally expanded with apex at
least four times wider than base 2
- Cercus elongate, more or less digitiform, with
apex not more than twice basal width 3
2. Antenna mostly orange-yellow with only the tip
of the first flagellomere dark brown; Till with
strong dorsal seta at %; cercus, expanded and
davate, subtriangular, and bearing distal row of
strong long pale setae, and with dorsal setose pro-
jection, (fig. 12) (Papua New Guinea)
A. maprik sp. n.
- Antenna dark brown; Till with short dorsal setae
on distal quarter; cercus, expanded and davate,
subtriangular, and bearing scattered pale setae (fig.
13) (Solomon Islands, New Guinea, Queensland)
A. nigricornis sp. n..
3. Cercus expanded, elongate and spatulate; sub-
epandrial sclerite broad with 3 strong apical setae
(fig. 19) (Micronesia: Truk) A. udot sp. n.
- Cercus not as above 4.
4. First flagellomere only weakly incised, if at all, so
that arista arises on the outer curvature of the first
flagellomere (fig. 15) 5
- First flagellomere deeply incised, so that arista
within distinct indentation on the outer curva-
ture of the first flagellomere (fig. 4) 7
5. Entire antenna yellow, with pale hairs; TI and Til
basally yellow, with distal three-quarters distinctly
white, and tarsomeres It,., white; basal epandrial
seta twice length of distal seta; cercus relatively
short and curved (fig. 14); (tropical Australia,
New Guinea, Solomon Islands, Vanuatu)
A. albitibia sp. n.
- At least first flagellomere partially brown; tibiae
and tarsi entirely yellow 6
6. Tibia III with long curved dorsal seta at % (fig.
10); antenna dark brown; cercus with strong
basal setae and apico-median field of short hairs
(Papua New Guinea) A. toma sp. n.
- Tibia III with short dorsal setae; scape and pedi-
cel yellow; first flagellomere mostly yellow with
only apex infuscated; cercus digitiform and rather
uniformly setose (fig. 6) (Papua New Guinea,
montane) A. putosa sp. n.
7. Antenna with at least scape and pedicel partially
yellow, and first flagellomere brown 8
- Antenna entirely dark brown 9
8. Cercus (fig. 1 1) lobate and elongate, almost twice
as long as epandrium, and with distinctive dorsal
digitiform projection which bears strong apical
seta; mesonotum dull brown-yellow; (New Gui-
nea, Solomon Islands) A. boharti sp. n.
- Cercus (fig. 1) short, about as long as epandrium,
and subtriangular; thorax dark brown/metallic
green (Solomon Islands, New Guinea, n. Aus-
tralia ) A. protractus Robinson
9. Epandrial lobe greatly prolonged and lobate,
completely covering surstylar arms, and subequal
to elongate digitiform cercus (fig. 8) (Solomon Is-
lands) A. malaita sp. n.
- Epandrial lobe relatively short, not much longer
than surstyli, and cerei subtriangular and tapering
10
10. Surstylus with dorsal arm strongly upcurved; cer-
cus elongate, about as long as epandrium (fig. 7)
(New Britain) A. colmanisp.n.
- Surstylus with arms subparallel; cercus lobate and
short, not extending much beyond apex of sur-
styli. (fig. 5) (Queensland) A. kurandasp. n.
Acropsilus protractus Robinson
(figs. 1-4, 20)
Acropsilus protractus Robinson 1963: 830. Holotype S ,
Solomon islands: Guadalcanal, Lunga River Valley,
16.ix.1944, J. L. Laffoon (usnm) [examined].
Additional material (all records based on males). - Aus-
tralia, northern territory: Berry Springs, 12.42'S
130.59'E, monsoonal rainforest, 15.vi.1954 (anic),
4.xii.l991-9.i.l992, 27.ix-30.x.l991, 30.x-4.xii.1991, 9.i-
28.ii.1992, malaise, 9-11.1.1992, yellow pans (ams); Bir-
raduk Ck, 17km WSW of Nimbuwah Rock, 12.19'S
133.13'E, 5.VÌ.1973; Cooper Ck, 11km SW of Nimbuwah
Rock, l.xi.1972; Kakadu NP, 19km NE of Baroalba Ck
Tijdschrift voor Entomologie, volume mi, 1998
Bickel: Australasian Acropsilus (Dolichopodidae)
Springs, 12.48'S 132.49'E, 28.x. 1972, malaise; East Alliga-
tor River, 5 km NNW of, 8-9-vi.l973 (anic); Koongarra, 16
km NE of Mt Cahill, 12.52'S 132.5 TE, 10.iii.1973; Lee
Point, 12.20'S 130.54'E, vi. 1964; Magela Ck, 2 km N of
Mudginbarry, 15.xi.1972; McArthur River, 2 km SSE of
Borroloola, 20.iv.1976; Surprise Ck, 45 km SSW of Bor-
roloola, 15.iv.1976 (anic). Kakadu NP, Gubara Springs
carpark, 17. i. 1992, pans (ams). - Queensland: Cockatoo
Creek, 11.39'S 142.27'E, 7-ii-2.iii.1993, 12.xi-l4.xii.1993,
malaise; Coen, 13.57'S 143.12'E, 16.viii-13.ix.1993, malai-
se; 1 km SE of Mt Cook, 15.30'S 145.16'E, 13.X.1980, ma-
laise; 3.5km SSW of Mt Baird, 17.26'S 145.04'E, 4.V.1981,
malaise; Bellenden Ker Range, 1km S of Cable Tower 6,
17.x-5.xi. 1981, malaise; Cedar Bay NP, Gap Ck, 5km ESE
of Mt. Finnigan, 15.50'S 145.20'E, 150 m, 14.V.1981, ma-
laise (anic), 14-I5.iv.1994, pans (ams); Davies Ck & Gold-
mine Ck, Kuranda, 3.V.1967, gen/sweep; Earl Hill, N of
Cairns, 16.48'S 145.42'E, 8.V.1967 (anic); Iron Range:
HkmENEofMtTozer, 12.43'S 143.18'E, ll-16.vii.1986,
malaise; 3km ENE of Mt Tozer, 12.45'S 143.14'E, 200 m,
28.vi-4.vii. 1986, malaise; 9 km ENE of Mt Tozer, 12.43'S
143.17'E, 5-10.VÜ-1986, malaise; Mcllwraith Range, 11km
NW of Bald Hill, 13.39'S 143.20'E, 520 m, rainforest,
26.vi-13.vii. 1989, malaise; Mossman Gorge, 16.26'S
145.16'E, 21-23.iv.1967; Mt Webb NP, 3 km NE of Mt
Webb, 15.03'S 145.09'E, 1. v. 1981, malaise; Rounded Hill,
15.17'S 145.13'E, 7.X.1980 (anic); Miriam Vale district, W
of Eurimbula, 24.20'S 151.34'E, 29.iii.1975; Mt Windsor
Tableland, Forestry Hut, 16.16'S 145.02'E, 1060 m,
16.iv.1994, pans; Half Ton Ck, NE of Mt Carbine, 16.29'S
145.13'E, 290 m, riverine forest, 18.iv.1994, pans; Water-
park Ck SF, nr Byfield, 50 m, rainforest, 2.XÜ.1992; Cathu
SF, O'Connell River, NW of Mackay, 30m, rainforest,
7.iv.l994, pans (ams). - western Australia: Carson Es-
carpment, Drysdale River, 14.49'S 126.49'E, 9-15.viii.1975,
malaise (anic). - Indonesia, irian jaya: Aru Islands, Tran-
gan, 1 km S of Popjetur, 06.48'S 134.04'E, 90 m, 23.vi-
1 1. vii. 1994, malaise (nmwc). - papua new guinea: Maprik,
21-22. iii.1964; Siutmeri, Sepik River, I6.iii.1964; Doa Es-
tate,80 km W of Port Moresby, 9.ix.l962; Eastern High-
lands South, Okapa, 27.x. 1964; Okapa, Kaga, 2,100 m,
11. vi. 1964; Finisterre Range, Damaindi, Madang Central,
1060 m, X.1964; Butemu III & VIII, Finisterre Range, 1270
m, x.1964 (anic); Normanby I., Wamula 29.xii.1988-
3.Ì.1989; Western Prov., Tabubil, 05.15'S 141.13'E, 650 m,
3.X.1993; Bainyik, 20.xii.1963 (ams); Bougainville: Boku,
4.vi.l956, & Ruisei, N of Tokinoitu, 2. vi. 1956 (bpbm). -
solomon islands: Rennell Island, Niupani, 22.viii.1962
(zmuc); Guadalcanal, v. 1944 (usnm); Velia Lavella, Are-
wana area, 17. xi. 1963; Florida Group, Nggela I., Haleta,
xi. 1975, 0-50m; Kolombangara Island: Ringgi Cove, xi.1976
& Kundulu, SW coast, 10. vii. 1959; Guadalcanal, Honiara,
200 m, xii.1975; Guadalcanal, Tadhimbko, 9.XÜ.1975;
Malaita, Auki, 100 m, xii.1975; Malaita, NW, Dala,
11. vi. 1964, light trap; New Georgia Group, Gizo I., 200 m,
xii.1975, ll.vii.1959; New Georgia, Munda, 15.vii.1959;
San Cristobal Island, Maniata, Kira Kira, xii.1975-i.1976;
Santa Ysabel, Buala, 26.vi.1960, light trap (bpbm).
Description
Male. - Length: 1.1; wing: 1.1X0.3.
Head. - Vertex and frons dark brown with metallic
green reflections and some grey pruinosity; face-
clypeus blue metallic green; palp and proboscis
brownish; scape and pedicel reddish-yellow to yellow,
although sometimes dorsally infuscated); first flagel-
lomere dark brown, rounded, setose, and deeply in-
cised.
Thorax. - Mesonotum dark brown with metallic
blue-green reflections, with some grey pruinosity; pleu-
ra matt brown; proepisternum with pale longer ventral
seta and 2 shorter dorsal setae near posterior margin.
Legs. - Coxae and remainder of legs yellow, al-
though CII sometimes infuscated; CI and CII with
short pale anterior hairs; CII and CIII each with short
brownish lateral seta; I: 2.0/ 1.5/ 0.7; 0.5; 0.4; 0.2;
0.2; leg I without distinctive setation; II: 2.0/ 2.2/ 0.8;
0.5; 0.3; 0.3; 0.2; Til with strong ad-pd pair at lA,
strong ad at 3/5 and apically, and with strong ventral
apical; III: 2.2; 2.1; 0.3/ 0.8/ 0.5/ 0.3/ 0.2; Till with
short ad-pd pair at Vi, and dorsal seta at %; Hit, some-
what globular and with a few ventral setae.
Wing. — Lower calypter yellow with fan of brown
setae; halter yellow.
Abdomen. - Entirely dark brown with sparse short
brown setae; hypopygium (fig. 1) brown, with dark
brown surstyli, and cream-white cerei; hypopygial
foramen left basolateral; hypandrium broad and api-
cally upcurved; aedeagus rather short and slightly
curved, and with notched phallus; ventral margin of
epandrium with stronger basal and shorter well-sepa-
rated epandrial setae; epandrial lobe with single apical
seta and externally overlapping digitiform surstylar
arms; cercus, expanded and davate, subtriangular,
and bearing long pale setae.
Female. - First flagellomere not as deeply incised;
oviscapt (figs 3, 4) divided into two hemitergites,
each with crest of 4 spine-like setae, and bearing elon-
gate digitiform process and stalked group of pedun-
culate setae.
Remarks
Acropsilus protr actus is widespread in northern Aus-
tralia, New Guinea and the Solomon Islands (fig. 20),
and is particular common in lowland habitats, but oc-
curs to elevations of 2000 m in Papua New Guinea.
In Australia, it is found across the monsoonal north,
and along the Queensland coast south to the Tropic
of Capricorn.
Figs. 1-6. Acropsilus species. - 1-4. Acropsilus protractus Robinson; 1, male postabdomen, left lateral; 2, female oviscapt, left
lateral; 3, female oviscapt, dorsal; 4, male antenna, left lateral. - 5-6. hypopygium and sternum 8, left lateral — 5, A. kuranda
sp. n.; 6, A. putosa sp. n. Abbreviations: aed, aedeagus; cer, cercus; epl, epandrial lobe; eps., epandrial seta; hyp, hypandrium;
sur, surstylus; st, sternum; sepsc, subepandrial sclerite; tg, tergum.
Tijdschrift voor Entomologie, volume mi, 1998
Acropsilus protractus has a yellow scape and pedicel
in contrast to the dark brown first flagellomere, al-
though in some specimens the dorsal scape and pedi-
cel are infuscated. The legs are entirely yellow. The
cerei appear somewhat apically truncated in dorsal
view. As well, there is some intraspecific variation in
cereal length, and specimens from northern Queens-
land appear to have slightly shorter cerei than Solo-
mon Island specimens.
Acropsilus kuranda sp. n.
(fig- 5)
Type material. - Holotype, S , Paratypes, $ , 2 9 ,
Australia. Queensland: 1 1 km along Black Moun-
tain, Kuranda State Forest, 20.iv.1967, D.H. Colless
(anic).
Additional material. Australia. Queensland: 6\ 29,
Bellenden Ker Nat. Park, Bartle Frere track, at junction of 3
creeks, 17.25'S 145.51'E, 200 m, rainforest, 12.iv.1994,
pans; â , Eungella NP, Crediton Creek, 920 m, rainforest,
30.xi.1992, pans; 3, 29, Little Cooper Creek, 16.1 O'S
145.27'E, 75 m, rainforest, l4.iv.1994; 3, Mason Ck, nr
Cape Tribulation, 16.05'S 145.28'E, 0-10 m, mangroves,
14-I5.iv.1994, pans; 3 , Mount Mirinjo, 18kmNWofIn-
nisfail, 25.iv.1980, sweep (ams); 23 , 9 , Mt Elliot NP, Alli-
gator Creek, 19.30'S 146.55'E, 30, sweep, 8.iv.l994 (cnc);
3 , 2 9 , Bramston Beach, nr Innisfail, rainforest, 30. iv. 1967;
3, Gillies Hwy, 3.5 km W of Little Mulgrave, 18.iv.1967;
2d, 9, Laceys Creek, near Mission Beach, 17.54'S
146.06'E, 13-14.V.1980; 3ò\ 11 9, Mcllwraith Range,
11km NW of Bald Hill, 13.39'S 143.20'E, 520 m, rainfor-
est, 26.vi-13.vii. 1989, malaise, 3ó\ The Boulders, NW of
Babinda, 17.22'S 145.55'E, 50 m, lO.v.1967, 8.vii.l967;
3 , 9 , Upper Mulgrave River, 1 6km from Goldsborough,
9.V.1967; 3, Wongabel SF, near Atherton, 17.19'S
145.31'E, 5. v. 1967 (anic).
Description
Male. - Length: 1.3; wing: 1.1X0.3.
Head. - Vertex, frons dark brown with metallic
blue reflections; face-clypeus metallic green; palp and
proboscis brown; antenna dark brown, first flagel-
lomere rounded, strongly setose, and deeply incised
(as in fig. 4), with apical arista arising from base of in-
cision.
Thorax. - Mesonotum dark brown with metallic
blue-green reflections, and with little pruinosity; pleu-
ra matt brown; proepisternum with longer ventral seta
and 2 shorter dorsal setae near posterior margin.
Legs. - Coxae and remainder of legs yellow; I: 1.8/
1.7/ 0.7; 0.5; 0.4; 0.3; 0.3; II: 2.3/ 2.5/ 0.8; 0.7; 0.3;
0.3; 0.2; Til with strong ad-pd pair at Vi, strong ad at
%, and with strong ventral apical; III: 2.6; 2.7; 0.3/
0.8/ 0.5/ 0.3/ 0.2; Till with short ad-pd pair at Va,
and a few short dorsal setae on distal quarter; lilt,
somewhat globular and with a few ventral setae.
Wing. - Lower calypter yellow with fan of brown
setae; halter yellow with dark brown club.
Abdomen. - Entitely dark brown with sparse short
brown setae; sternum 8 with setae as figured; hypopy-
gium (fig. 5) brown with dark brown surstyli and
cream-white cerei; hypopygial foramen left basolater-
al; hypandrium broad and apically upcurved; aedea-
gus rather short and slightly curved, and with
notched phallus; ventral margin of epandrium with
stronger basal and shorter distal epandrial setae; epan-
drial lobe with 2-3 apical setae, and externally over-
lapping subtriangular surstylar arms; subepandrial
sclerite Y-shaped with two apical arms, each ending in
strong seta; cercus lobate, bearing long pale setae, and
not extending much beyond apex of surstyli.
Female. - Similar to male except as noted: eyes
slightly more widely separated, and facets of uniform
size; first flagellomere not as deeply incised; Hit,
somewhat less swollen.
Remarks
Acropsilus kuranda occurs in tropical forests of
northeastern Queensland, from the Mcllwraith Range
of Cape York Peninsula south to Eungella, west of
MacKay. Most specimens were taken in the Cairns
disttict. This species is close to the sympatric A. pro-
tractus, but differs in having the antenna entirely dark
brown, a short lobate cercus which doesn't extend
much beyond the sutstyli, and a more deeply incised
first flagellomere.
Acropsilus colmani sp. n.
(fig- 7)
Type material. - Holotype, 6 , Paratypes 5 ó* 2 ? ,
Papua new guinea, New Britain, Amelei Village,
Fullerborn Harbour, xii.1988, P H Coiman (ams).
Additional material. - papua new guinea, 3 , 9 , New
Britain, Keravat, Rabaul, 100 m, 15-3 1.x. 1968, N L H
Krauss (bpbm); 3, Kandanggei, Sepik River , 2.iii.l964
(anic).
Description
Male. - Length: 1.2-1.3; wing: 1.1X0.4.
Head. - Vettex, frons dark brown with metallic
gteen reflections clypeus and face bright metallic
green; palp and proboscis brown; antenna uniformly
dark brown, first flagellomere rounded, setose, but
only shallowly incised.
Thorax. - Mesonotum dark brown with metallic
blue-green reflections, and shining, with little pru-
inosity; pleura matt brown; proepisternum with
longer ventral seta and 2 shorter dorsal setae near pos-
terior margin.
Legs. - Coxae and remainder of legs yellow; Til
with strong ad-pd pair at Va, strong ad at 3/5, and with
strong ventral apical; Till with short ad-pd pair at lA,
Bickel: Australasian Acropsilus (Dolichopodidae)
Figs. 7-10. Acropsilus species. - 7-9, hypopygium and sternum 8, left lateral; 10, male tibia and tarsus III, anterior- - 7 Acrop
silus colmarli sp. n.; 8, A. malaita sp. n.; 9-10, A. toma sp. n. anterior, /, Acrop
Tijdschrift voor Entomologie, volume i4i, 1998
and a few short dorsal setae on distal quarter; Hit,
somewhat globular and with a few ventral setae.
Wing. - Lower calypter yellow with fan of brown
setae; halter yellow with brownish club.
Abdomen. - Preabdomen dark brown with sparse
short brown setae, postabdomen mostly light brown
except hypopygium with dark brown surstyli and
elongate cream-white cerei; hypopygium (fig. 7); hy-
pandrium broad and upcurved apically; aedeagus
curved, and with notched phallus; ventral margin of
epandrium with well-separated strong basal and
shorter epandrial setae; epandrial lobe with short api-
cal setae, and externally overlapping surstylus; sursty-
lus dark brown, with dorsal arm strongly upcurved;
subepandrial sclerite cercus with 4 strong apical setae,
each on distinct peduncle; elongate and oblong, and
bearing pale setae.
Female. - Similar except lack mssc and as noted:
abdomen entirely dark brown.
Remarks
Acropsilus colmarti is known from New Britain and
mainland Papua New Guinea. This species is close to
A. protractus and also has a hairy, incised first flagel-
lomere. However, the combination of entirely dark
brown antenna, elongate cercus, and upcurved sursty-
lar arm are diagnostic. This species is named for Phil
Colman, who has collected extensively in Melanesia.
Acropsilus malaita sp. n.
(fig. 8)
Type material. - Holotype, <$ , Solomon Islands:
Malaita (NW), Dala, 5.VÌ.1964, light trap, R. Straat-
man (bpbm).
Description
Male. - Length: 1.3; wing: 1.2X0.3.
Head. - Vertex and frons dark brown with metallic
reflections; face clypeus metallic green; antenna dark
brown, first flagellomere rounded, weakly setose, and
only slightly incised.
Legs. — Coxae and remainder of legs yellow; Til
with strong ad-pd pair at Va, strong ad at 3/s, and with
strong ventral apical; Till with short ad-pd pair at Va,
and short dorsal setae near %; It! somewhat globular
and with ventral setae.
Wing. - Lower calypter yellow with fan of brown
setae; halter yellow with infuscated club.
Abdomen. - Entirely dark brown with sparse short
brown setae; hypopygium (fig. 8) with dark brown
surstyli, and cream-white cerei; hypandrium broad
and apically upcurved; aedeagus rather short and
slightly curved; ventral margin of epandrium with
very strong basal epandrial seta which is somewhat
laterally displaced and shorter distal epandrial seta;
epandrial lobe dark brown, greatly expanded and dis-
tally prolonged with some short apical setae, and to-
tally covering the more median surstylar arms, one of
which is distally bent; subepandrial sclerite cercus
with 4 strong apical setae, each on distinct peduncle;
cercus elongate and digitiform.
Female. - Unknown.
Remarks
Acropsilus malaita is known only from the Malaita,
Solomon Islands type locality. The greatly prolonged
and expanded epandrial lobe and elongate digitiform
cercus are diagnostic.
Acropsilus toma sp. n.
(figs. 9-10)
Type material. - Holotype, S , Paratypes 9 , papua
new guinea: Mangalese area, nr Toma, SSW of Po-
pendetta, 600 m, ix.1964, R. Pullen (anic).
Additional material. - papua new guinea: ó\ Central
Prov., Port Moresby, 29.V.1984 (cmnh).
Description
Male. - Length: 1.3; wing: 1.1X0.3.
Head. - Vertex, frons, face, clypeus dull dark
brown; palp and proboscis brown; antenna dark
brown, first flagellomere rounded, not densely haired
and only weakly incised, with apical arista arising
from base of incision.
Thorax. - Mesonotum dark brown with some
brown pruinosity; pleura matt brown; setae black.
Legs. - Coxae brown but yellowish distally; re-
mainder of legs yellow, although femora somewhat
infuscated; Til with strong ad-pd pair at Va, strong ad
at %, and with strong ventral apical; Till (fig. 10)
with short ad seta at Vs and with very long curved
dorsal seta at % (mssc); Hit, somewhat globular and
with some ventral setae.
Wing. - Lower calypter yellow with fan of brown
setae; halter brownish with dark brown club.
Abdomen. - Entirely dark brown with sparse short
brown setae; hypopygium (fig. 9) brown with dark
brown surstyli and white cerei; hypandrium broad
and apically upcurved; aedeagus curved, and with
notched phallus; ventral margin of epandrium with
stronger basal and slightly shorter distal epandrial se-
tae; epandrial lobe externally overlapping surstylar
arms; subepandrial sclerite curved and elongate, with
apical setae; cercus oblong, bearing row of long pale
ventral setae and field of strong curved basomedial
seat, and with apicomedial field of short hairs.
Female. - Similar to male except as noted: eyes
slightly more widely separated, and facets of uniform
size; Till with only short dorsal seta at %.
10
BicKEL: Australasian Acropsilus (Dolichopodidae)
Figs. 11-13, Acropsilus species, hypopygium and sternum 8, left lateral. -11, Acropsilus bohartisp. n.; 12, A. mapriksp. n.; 13,
A. nigricornis sp. n.
11
Tijdschrift voor Entomologie, volume hi, 1998
Remarks
Acropsilus toma is known from two lowland sites in
Papua New Guinea. It is distinguished by the long
curved dorsal seta of male tibia III (mssc), and the
row of long setae along the ventral border of the ob-
long cerei.
Acropsilus boharti sp. n.
(fig. 11)
Type material. - Holotype, S , Paratypes, â , 29 ,
Indonesia. Irian JAYA, Biak I., 25.iv.1945, G.E. Bo-
hart (cas).
Additional material. - solomon islands: o\ Guadal-
canal, Tambalia, 30 km W of Honiara, malaise trap,
21.v.1964(bpbm).
Description
Male. - Length: 1.2; wing: 1.3X0.4.
Head. — Vertex, frons dark brown with metallic re-
flections; face-clypeus metallic blue-green; palp and
proboscis yellowish; antenna yellowish with first fla-
gellomere slightly infuscated; first flagellomere
rounded, strongly setose and incised, with apical
arista arising from base of incision.
Thorax. — Mesonotum dull brown-yellow with lit-
tle pruinosity; pleura yellow.
Legs. - Coxae and remainder of legs yellow; Til
with ad-pd pair at Va, strong ad at %, and with strong
ventral apical; Till with short ad-pd pair at Va, and
strong subapical dorsal seta; lilt, somewhat globular
and some ventral setae.
Wing. - Lower calypter yellow with fan of brown
setae; halter yellow with dark brown club.
Abdomen. - Entirely dark brown with sparse short
brown setae; hypopygium (fig. 11) brown with dark
brown surstyli and white cerei; hypandrium broad
and apically upcurved; aedeagus curved with indent-
ed phallus; ventral margin of epandrium with strong
basal and distal epandrial setae; epandrial lobe with
short setae and externally overlapping rather short
surstylar arms; subepandrial sclerite with group of 3
strong setae, each arising from separate long pedun-
cle; cercus elongate and lobate, bearing pale setae, and
with distinctive dorsal digitiform projection which
bears strong apical seta.
Female. — Similar to male except as noted: eyes
slightly more widely separated, and facets of uniform
size; first flagellomere not as deeply incised; Hit,
somewhat less swollen.
Remarks
Acropsilus boharti is known from two lowland Me-
lanesian sites: Biak Island off northern Irian Jaya, and
Guadalcanal, Solomon Islands, but it probably occurs
in intervening areas. This species is distinguished by
the digitiform projection along the dorsal surface of
the elongate cercus. This digitiform projection is
longer on the Irian Jaya males (figured) than on the
Solomon Island male.
Acropsilus maprik sp. n.
(fig. 12)
Type material. - Holotype, S , papua new guinea:
Maprik, 21.iii.1964, D.H. Colless (anic).
Description
Male. - Length: 0.9; wing: 1.2X0.4.
Head. - Vertex and frons dark brown, with metallic
green reflections; face-clypeus metallic green-bronze;
palp and proboscis brown; antenna mostly orange-
yellow with only the tip of the first flagellomere dark
brown; first flagellomere rounded, setose, and only
shallowly incised.
Thorax. — Dark brown with metallic blue-green re-
flections, and shining, with little pruinosity; pleura
matt brown.
Legs. - Coxae and remainder of legs yellow, al-
though FII and Fill slightly infuscated; Til with
strong ad-pd pair at Va, strong ad at 3A, and with
strong ventral apical seta; Till with short ad-pd pair
at Va, and strong dorsal seta at %; Hit, with ventral roe
of short hairs (mssc).
Wing. — Lower calypter yellow with fan of brown
setae; halter yellow.
Abdomen. - Preabdomen dark brown with sparse
short brown setae; postabdomen and hypopygium
(fig. 1 2) mostly yellow, except hypopygium with dark
brown surstyli and ivory-white cerei; hypandrium
broad and apically upcurved; aedeagus rather short
and slightly curved; ventral margin of epandrium
with stronger basal and shorter epandrial setae; epan-
drial lobe with single apical seta and externally over-
lapping digitiform surstylar arms; subepandrial scle-
rite with 4 strong, slightly pedunculate setae; cercus,
expanded and davate, subtriangular, and bearing dis-
tal row of strong long pale setae, and with medio-dor-
sal setose projection.
Female. — Unknown.
Remarks
Acropsilus maprik is known only from northern
lowland Papua New Guinea. The enlarged davate
cercus with its distal row of strong setae and yellow
antennae are diagnostic. This species is close to A. ni-
gricornis and both species have davate cerei.
12
Bickel: Australasian Acropsilus (Dolichopodidae)
Acropsilus nigricornis sp. n.
(fig. 13)
Type material. - Holotype, 6 , solomon islands.
New Georgia Group, Gizo L, 0-100 m, ii.1984, N. L.
H. Krauss (bpbm).
Additional material. - Australia. Queensland: 3 , Iron
Range, nr. Mt Lamond, 20.xii.1971 (ams); o\ Mt Webb
NP, 3 km NE of Mt Webb, 15.03'S 145.09'E, 2.X.1980,
malaise trap (anic). - Indonesia, irian jaya: o*, Biak I.,
25.iv.1945 (cas).
Description
Male. - Length: 1.1; wing: 1.0X0.3.
Head. - Vertex, ftons, face-clypeus metallic green,
dark brown, shining with metallic blue reflections;
palp and proboscis brown; antenna dark brown, first
flagellomere rounded, setose, and weakly incised.
Thorax. - Mesonotum dark brown with metallic
blue-green reflections, and with little pruinosity; pleu-
ra matt brown; proepisternum with longer ventral seta
and 2 shorter dorsal setae near posterior margin.
Legs. - Coxae and remainder of legs yellow; I: 2.0/
1.5/ 0.7; 0.3; 0.3; 0.2; 0.2; II: 2.0/ 2.2/ 0.8; 0.5; 0.3;
0.3; 0.2; Til with strong ad-pd pair at lA, strong ad at
3/5, and with strong ventral apical; III: 2.2; 2.1; 0.3/
0.8/ 0.5/ 0.3/ 0.2; Till with short ad-pd pair at Va,
and a few short dorsal setae on distal quarter; lilt,
somewhat globular and with a few ventral setae.
Wing. - Lower calypter yellow with fan of brown
setae; halter yellow.
Abdomen. - Entirely dark brown with sparse short
brown setae; hypopygium (fig. 13) brown, with dark
brown surstyli, and cream-white cerei; hypandrium
broad and apically upcurved; aedeagus rather short
and slightly curved; ventral margin of epandrium
with well- separated stronger basal and shorter epan-
drial setae; epandrial lobe with single apical seta and
externally overlapping digitiform surstylar arms;
subepandrial sclerite with distal curved Y-shaped
arms; cercus, expanded and davate, subtriangular,
and bearing scattered long pale setae.
Female. - Unassociated.
Remarks
Acropsilus nigricornis is known from the Solomon
Islands, Cape York Peninsula, and offshore northern
Irian Jaya. The enlarged davate cercus, dark brown,
weakly-incised first flagellomere, and shining metallic
blue face are diagnostic.
Acropsilus albitibia sp. n.
(figs. 14-18,21)
Type material. - Holotype â , Parartypes 4 6 , 3 ? ,
Australia. Queensland: Iron Range, 3.2 km NE of
Mt Lamond, 22.xii.1971, D.K. McAlpine & G.A.
Holloway. Paratypes, 8 cT, 21 9, 1.6 km W of Mt La-
mond, 13-23.xii.1971, malaise; AS , 29, Claudie Riv-
er, nr Mt Lamond, 12.48'S l43.13'E,3.vi. 1966 (ams).
Additional material (all records based on males). - Aus-
tralia, northern territory: Berry Springs, 12.42'S
130.59'E, monsoonal rainforest, 30.x-4.xii. 1991, 4.xii. 1991-
9.Ì.1992, malaise (ams); Darwin, 12.27'S 130.50'E, vi-
vii.1958, at light (anic); Kakadu NP, Baroalba Ck Springs,
19km NE of Mt. Canili, 12.48'S 132.49'E, 29.X.1972,
malaise & I6.xi.1972, at light (anic); Nourlangie Ck, 8 km
N of Mt Cahill, 26.X.1972; Lee Point, 12.20'S 130.54'E,
vi. 1964 (anic); Magela Ck, 2 km N of Mudginbarry,
15.xi.1972 (bpbm). - Queensland: 14 km NW of Hope
Vale Mission, 15.16'S 144.49'E, 8.X.1980, malaise; 1 km
SE of Mt Cook, 15.30'S 145.16'E, 13.X.1980, malaise; 4
km SSE of Cape Tribulation, 16.06'S 145.29'E, 21-
22.xi.1981; 5 km NW of Rounded Hill, 15.17'S 145.10'E,
7.X.1980, malaise; Bamboo Ck, nr Mail, 16.19'S 145.23'E,
25.iv.1967; Cedar Bay NP, Gap Ck, 5 km ESE of Mt
Finnegan, 15.50'S 145.20'E, 150 m, 15.V.1981, malaise;
Earl Hill, N of Cairns, 16.48'S 145.42'E, 8.V.1967; Gillies
Highway, 3 km W of Little Mulgrave, 18.iv.1967; Iron
Range, 3 km ENE of Mt Tozer, 12.45'S 143.14'E, 200 m,
28.vi-4.vii. 1986, malaise; 9 km ENE of Mt Tozer, 12.43'S
143.17'E, 5-10.vii.1986, malaise; 9 km NW of Mt Tozer,
30-vi-7.vii.1986; Laceys Ck, nr Mission Beach, 17.54'S
146.06'E, 13-14.V.1980; Station Ck, 7 km WSW of Hope
Vale Mission, 15.19'S 145.03'E, lO.v.1981; The Boulders,
NWof Babinda, 17.22'S 145.55'E, 50 m, 8.VÜ.1971; Mcll-
wraith Range, 11km NW of Bald Hill, 13.39'S 143.20'E,
520m, rainforest, 26.vi-13.vii.1989, malaise; Mt Cook NP,
15.29'S 145.16'E, lO.v.1981, 12.x. 1980, malaise; Mt Webb
NP, 3 km NE of Mt Webb, 15.03'S 145.09'E, 1-2.V.1981,
malaise (anic); North Maria Ck, nr Silkwood, 17.45'S
146.02'E, I4.xii.1961; Double Mouth Ck, 30 km NE of
Heathlands, 11.37'S 142.49'E, 22.iii.1992; Portland Roads,
12.36'S 143.25'E, I4.xii.1971, malaise; Silkwood, 17.45'S
146.01'E, 25.V.1958; Tully River, 17.46'S 145.36'E, 175
m, rainforest, 10.iv.1994, pans; Bertie Ck pump, nr Heath-
lands, 11.46'S 142. 36'E, 21. iii. 1992, malaise (ams). -west-
ern Australia: Kimbolton, xi-xii.1982, malaise (anic). —
Indonesia: Aru Islands, Trangan, 1 km S of Popjetur,
06.48'S 134.04'E, 90 m, 23.vi-ll.vii.1994, malaise trap
(nmwc); - irian jaya: Biak I., 25.iv.1945 (cas); Oransbari,
28. vii. 1962 (bpbm). - papua new guinea. Central Prov.:
Doa Estate, 80 km W of Port Moresby, 2-9.ix.1962 (anic);
5 km NW of Brown River Bridge, 6.ix.l984; Aroa River,
Aroana Estate, 2.XÜ.1963, 26.viii.1984; Imbia, nr Maprik,
19.xii.1963 (ams); Rouna, 300 m, xi.1968; Brown River,
dry riverbed, 30.viii. 1959 (bpbm'. - solomon islands.
Guadalcanal, Honiara, 200 m, xii.1975; Guadalcanal, Poha
River, 5 m, 2.VÜ.1959; Malaita, Auki, 100 m, 18.xi.1957;
New Georgia Group, Ghizo I., 200 m, xii.1976, malaise;
Rendova Island, Agagana, 13.xi.1970; Dai Island, Bethle-
hem, 10 m, xii.1972 (bpbm). - Vanuatu. Lopevi, Lamen,
100 m, ii.1976 (bpbm).
Description
Male. - Length: 1.2; wing 1 . 1 X 0.4
Head. - Vertex and frons dark brown with metallic
green reflections; face and clypeus bright metallic
green; palp and proboscis brown; antenna entirely red-
13
Tijdschrift voor Entomologie, volume i4i, 1998
Figs. 14-19. Acropsilus species. - 14-18. Acropsilus albitibia sp. n.; 14, male postabdomen, left lateral; 15, male head, anteri-
or; 16, male wing, dorsal; 17, male thorax and abdomen, posterodorsal; 18, internal mouthparts, left lateral; 19, A. udot sp. n.,
hypopygium and sternum 8, left lateral.
14
Bickel: Australasian Acropsilus (Dolichopodidae)
Acropsilus protractus
Fig. 20. - Distribution, Acropsilus protractus Robinson.
dish-yellow; first flagellomere rounded, globular, with
pale apical hairs, but without apical notch (fig. 15).
Thorax. - Brownish with metallic green reflec-
tions, with some brown pruinosity over mesonotum
and pleura, and with humeral area yellowish; proepis-
ternum with 3 short spaced setae near posterior mar-
gin; setae black.
Legs. — Coxae and femora yellow; TI and Til basal-
ly yellow, with distal three-quarters distinctly white
or ivory coloured (mssc), and especially visible in an-
terior view, strongly contrasting with yellow ground
colour; Till pale yellow; tarsi mostly yellow, but It23
white (mssc); CI and CII with short pale anterior
hairs; CIII and CIII each with short brown lateral
seta; I: 2.7/ 2.2/ 0.8; 0.4; 0.3; 0.4; 0.4; leg I without
distinctive setation; II: 3.6/ 3.2/ 1.0; 0.8; 0.4; 0.4;
0.4; FII with short subapical av and pv setae; Til with
strong ad-pd pair at lA, strong anterior at and apical-
ly, and with strong apico-ventral seta; III: 3.0; 3.4;
0.4/ 1.2/ 0.6/ 0.5/ 0.4; Fill with short subapical av
and pv setae; Till with strong ad and pd at V4, and
with strong subapical dorsal seta; Hit, somewhat
globular with short ventral setae.
Wing (fig. 16). - Lower calypter yellow with fan of
brown; halter yellow.
Abdomen. - Preabdomen (fig. 17) dark brown with
short yellowish setae; postabdomen (segments 6-9)
and hypopygium yellowish, except surstyli dark
brown and cerei ivory white; hypopygium (fig. 14);
hypandrium broad and hood-like; ventral margin of
epandrium bearing very strong basal epandrial seta
and shorter distal seta; epandrial lobe digitiform with
apical and subapical setae; surstylar arms slightly
curved and digitiform; subepandrial sclerite elongate,
with basal and distal pedunculate setae; cercus subtri-
angular and tapering and covered with white setae.
Female. - Similar to male except lack mssc and as
noted: face wider; TI and Til entirely yellow; ab-
domen dark brown.
Remarks
Acropsilus albitibia is found in monsoonal northern
Australia, from Cape York Peninsula to the Kimber-
ley district, Western Australia, and widely across low-
land Melanesia, from the Aru Islands and mainland
New Guinea to the Solomon Islands and Vanuatu
(fig. 21).
The ivory-white colour on male tibiae I and II is di-
agnostic. In addition, most males have similar white
tarsomeres 2-3 on leg I, but this is sometimes indis-
tinct, so that tarsus I appears entirely pale yellow. Also
it should be noted that in some specimens the white
coloration on tibiae I and II is only weakly developed.
This might be due to age, state of preservation, or in-
traspecific variation. Of particular interest, the ivory-
white colour on both male legs I and II is identical to
the male cereal colour (and the cereal colour of most
Acropsilus, species), and possibly both the legs and cer-
ei are used in courtship display.
15
Tijdschrift voor Entomologie, volume i4i, 1998
\
\^1
•^>
H
Tropic of Capricorn
Fig. 21.- Distribution, Acropsilus albitibia sp. n.
Acropsilus putosa sp. n.
(fig. 6)
Type material. - Holotype 6, Paratype 6 papua
new guinea. Eastern Highlands, Putosa, 2400 m (as
8000 ft.), vii. 1968, R. Hornabrook (antic).
Description.
Male. - Length: 1.3; wing: 1.6X0.5.
Head. — Vertex and ftons dark brown with metallic
green reflections and some grey pruinosity; face-
clypeus blue metallic green; palp and proboscis brown-
ish; scape and pedicel yellow; first flagellomere mostly
yellow with only apex infuscated, and rounded, not
incised, and weakly haired.
Thorax. - Mesonotum dark brown with metallic
blue-green reflections, with some grey pruinosity;
pleura matt brown; proepisternum in basal third with
longer ventral seta and shorter dorsal seta near poste-
rior margin.
Legs. - Coxae yellow, although CII with some basal
infuscation; femora mostly brown, and remainder of
legs yellow, Til with strong ad-pd pair at Va, strong ad
at Ys, and with strong ventral apical; Till with short
ad-pd pair at Va, and short dorsal setae at %; lilt,
somewhat globular and with a few ventral setae.
Wing. - Lower calypter yellow with fan of brown
setae; halter yellow.
Abdomen. - Entirely dark brown with sparse short
brown setae; hypopygium (fig. 6) brown, with dark
brown surstyli, and cream-white cerei; hypandrium
broad and apically upcurved; aedeagus upcurved; ven-
tral margin of epandrium with subequal basal and dis-
tal epandrial setae; epandrial lobe with group of short
apical setae; surstylar arms overlapping and digiti-
form; cercus elongate, digitiform and with pale setae.
Female. - unknown.
Remarks
Acropsilus putosa is known only from the eastern
highlands of Papua New Guinea. The elongate digit-
form cercus is diagnostic. The antenna is mostly yel-
low, with the first flagellomere weakly incised, and
sparsely haired, and this species is close to A. albitibia.
Acropsilus udot sp. n.
(fig. 19)
Type material. - Holotype, S federated states
of micronesia: Truk Group: Udot I, 25.V.1946,
H.K. Townes (usnm).
Additional material. - federated states of microne-
sia: ô , Truk Group: Moen I., Mt. Teroken N, 28.xii.1952
(bpbm).
Description
Male. - Length: 1.3; wing 0.9X0.3.
Head. - Vertex, frons dark brown with metallic
green reflections; face and clypeus metallic green;
palp and proboscis yellow; scape and pedicel entirely
yellow; first flagellomere and arista missing.
Thorax. - Mesonotum brown with little pruinosi-
16
Bickel: Australasian Acropsilus (Dolichopodidae)
ty; pleura matt brown.
Legs. — Coxae and remainder of legs yellow; Til
with strong ad-pd pair at Va, strong ad at 3/5, and with
strong ventral apical; Till with short ad-pd pair at Va,
and short dorsal setae at %; Hit, globular and with
some ventral setae.
Wing. - Lower calypter yellow with fan of brown
setae; halter yellow.
Abdomen. - Dark brown with sparse short brown
setae; hypopygium (fig. 19) brown, with dark brown
surstyli, and cream-white cerei; ventral margin of
epandrium with stronger basal and shorter distal
epandrial setae; epandrial lobe with two apical setae,
and externally overlapping digitiform surstylar arms;
subepandrial sclerite broad with 3 strong, spaced api-
cal setae; cercus expanded, elongate and spatulate,
bearing weak setae.
Female. - Unknown.
Remarks
Acropsilus udot is known only from the Truk
Group in the Caroline Islands. The elongate spatulate
cercus is diagnostic. Although both specimens are
missing the first flagellomere, based on the scape and
pedicel, this species seems close to A. albitibia.
References
Becker, T., 1918. Dipterologische Studien. Dolichopodi-
dae. Dritter Teil. - Nova Acta Academiae Caesareae
Leopoldino Carolinae 104: 35-214.
Bickel, D. J. & C. E. Dyte, 1989. Family Dolichopodidae.
- In: N. Evenhuis (ed.), Catalog of Australasian and Oce-
anian Diptera, pp. 393-41 8. Bishop Museum Press, Hon-
olulu.
Bickel, D. J., 1995. Insects of Micronesia. Volume 13, no.
8. Diptera: Dolichopodidae Part I. Sciapodinae, Medete-
rinae and Sympycninae (part). - Micronesica 27, 73-1 18.
Couturier, G., 1977. Les Diptères Dolichopodidae de Côte
d'Ivoire: description de trois nouvelles espèces. - Bulletin
de la Société entomologique de France 82: 220-225.
Cumming, J. M., B. J. Sinclair, & D. M. Wood, 1995. Ho-
mology and phylogenetic implications of male genitalia
in Diptera - Eremoneura. - Entomologica Scandinavica
26: 120-152.
Grootaert, P. & H. J. G. Meuffels, 1997. Griphomyia (Di-
ptera, Dolichopodidae, Peloropeodinae), a new genus from
Thailand. - Belgian lournal of Zoology 127: 107-1 14.
Hollis, D., 1964. Notes and descriptions of Indonesian
Dolichopodidae in the Zoologisch Museum, Amsterdam.
- Beaufortia 10: 239-274.
Lamb, C. G., 1922. The Percy Sladen Trust expedition to
the Indian Ocean in 1905, under the leadership of Mr. J.
Stanley Gardiner, M.A. Vol. 7. No. Vili. Diptera: Asili-
dae, Scenopinidae, Dolichopodidae, Pipunculidae, and
Syrphidae. - Transactions of the Linnean Society of Lon-
don (2, Zoology) 18: 361-416.
Loew, H., 1869. Beschreibung europäischer Dipteren I. Sys-
tematische Beschreibung der bekannten europäischen
zweiflügeligen Insecten von Johann Wilhelm Meigen.
Neunter Theil. - H. W. Schmidt, Halle. 319 pp.
Lundbeck, W., 1912. Diptera Danica, Part IV: Dolichopo-
didae. - S. Gad, Copenhagen. 416 pp.
McAlpine, J. F., 1981. Morphology and Terminology
Adults. - In: J. F. McAlpine et al., Manual of Nearctic
Diptera, Vol. 1, Research Branch Agriculture Canada,
Ottawa, Monograph 27: 9-64.
Mik, J., 1878. Dipterologische Untersuchungen. -Jahres-
berichte der. Kaiserlich-königlichen Akademie, Gymnasi-
um, Wien 1878: 1-24.
Negrobov, O. P., 1984. The genera of the family Dolicho-
podidae Diptera new for the faunas of the Palearctic and
USSR. - Zoologicheskii Zhurnal 63: 1 1 1 1-1 1 15. [in
Russian] .
Negrobov, O. P., 1986. On the system and phylogeny of flies
of the Fam. Dolichopodidae (Diptera). - Entomologich-
eskoye Obozreniye 65: 182-186 [English translation, En-
tomological Review, Washington (1987) 65: 16-20].
Parent, O., 1938. Diptères Dolichopodidae. - Faune de
France 35. Paris, 720 pp.
Robinson, H., 1963. A new species of Acropsilus from the
Solomon Islands (Diptera: Dolichopodidae). - Canadian
Entomologist 95: 830-831.
Robinson, H. & J. R. Vockeroth, 1981. Dolichopodidae. -
In: J. F. McAlpine et al., Manual of Nearctic Diptera Vol
1. - Research Branch Agriculture Canada, Ottawa,
Monograph 27: 625-639.
Ulrich, H., 1981. Zur systematischen Gliederung der
Dolichopodiden (Diptera). - Bonner Zoologische
Beiträge 31: 385-402.
Received: 29 May 1998
Accepted: 23 June 1998
17
Tijdschrift voor Entomologie, volume ui, 199s
BOOK REVIEWS
Zlata S. Gershenson & Sandrine A. Ulenberg, 1998. The
Yponomeutinae (Lepidoptera) of the World exclusive of
the Americas. - Koninklijke Nederlandse Akademie van
Wetenschappen, Verhandelingen Afdeling Natuur-
kunde, tweede Reeks, Vol. 99. Amsterdam, etc. 250 pp,
many figs., 3 colour plates, hardback 25x17,5 cm. [ISBN
0-444-85819-9]. Price NLG 125.00.
This books presents an overview of the Yponomeu-
tinae of the old world, with 233 species, including
fossil ones. All the species are listed alphabetically
with references, type deposition and information on
biology (flight period, hostplants). Some 22 less
known species are fully described, with genitalia fig-
ures. One new genus and eight new species are de-
scribed, nine species are transferred to other genera
and five new synonyms are established. Three colour
plates show 62 species in natural size.
Introductary chapters contain a.o. a historical re-
view, a list of hostplants, a checklist and a list of Ypo-
nomeutid genera excluded from this subfamily. Also a
key for the 25 listed genera and a key for the species
of Yponomeuta is provided.
This book is an important basis for any further tax-
onomie and biological work on this highly interesting
group of micro-moths. Especially the supposed an-
cient relationship with the hostplant family Celas-
traceae is intriguing. In this light it is to be regretted
that the book is not completed with the American
species. It is true that even for a 'simple' checklist ma-
terial ought to be checked, but a mere list, even not
checked, of names would have been helpful.
Strong points of the book are the completeness of
references in the annotated list, the type-data and
hostplant-data, the lists of hostplants and the list of
excluded genera.
Some points of criticism are: the absence of gener-
ic diagnoses makes it impossible to judge why species
are placed in a certain genus; many of the pho-
tographed moths are too small, some are hardly rec-
ognizable. I personally do not like long citations of
references in the check-list and would have preferred
short references and a list of full references.
Despite this, the book is recommended for Lepi-
doptera taxonomists and evolutionists working with
this group of insects. I hope the authors will now find
time and endurance to continue with the next step: a
phylogenetic analysis of this subfamily.
Menno Schilthuizen & Henk Vallenduuk, 1998. Kevers op
kadavers. - Wetenschappelijke Mededeling KNNV, 222,
148 pp. [ISBN 90-5011-112-2]. Price NLG 29,50 excl.
p&p. [Beetles on carrion, In Dutch]
This booklet gives an overview of the Dutch bee-
tles, living on carrion. It includes an identification
key and descriptions of 22 species of Histeridae, 16
species of Silphidae and 23 Cholevidae. For these
species also distribution maps are provided. For an-
other 10 beetle families short keys are given for car-
rion feeding genera, or only a short description of the
family.
A nice booklet for all entomologists interested in
beetles or carrion fauna; also of interest for forensic
entomologists.
Johan van Zoest (ed.), 1998. Biodiversiteit. - KNNV Uitge-
verij, Utrecht, 212 pp. Hardback 17x25 cm. [ISBN 90-
5011-107-6]. Price NLG 59,50 excl. p&p. [Biodiversity,
In Dutch]
This book is a general introduction into the subject
of Biodiversity for the interested public in The Ne-
therlands. It deals with the various aspects of Biodiver-
sity and has special chapters on ecological background,
the natural and artificial landscapes, the vulnerability
of species for extinction, the social-economic side of
biodiversity and a chapter with the intriguing title
'Single ticket to the bare lands? Rather not.'
The book is well written, and probably will help
the public to understand the problems of biodiversi-
ty; but reathers not familiar with Dutch will not find
much in the book; even a summary is missing.
It is curious further that amongst the authors are
hardly real specialists of the biodiversity: taxonom-
nists, particularly those of arthropods. May be this is
the explanation that some recent publications, even
from the same publisher, were overlooked, or at least
not cited. Such as the previous book on Biodiversity
in The Netherlands (1995) and the recent Ortho-
ptera book (see review on page 48). Instead, distribu-
tion data of Orthoptera and Odonata were taken
from much older sources, partly out of date. The pho-
tograph depicting Tettigonia viridissima shows a very
young larva, but doesn't mention this, so that laymen
probably mistake this for an adult.
The book is only recommended as an introduc-
tion, for those familiar with Dutch.
[Erik J. van Nieukerken]
18
SEI-WOONG CHOI
Finnish Museum of Natural History, Helsinki
SYSTEMATICS OF THE GENUS HETEROTHERA
INOUE (LEPIDOPTERA, GEOMETRIDAE:
LARENTIINAE)
S.-W. Choi, 1998, Systematics of the genus Heterothera Inoue (Lepidoptera, Geomecridae: Lar-
entiinae). - Tijdschrift voor Entomologie 141: 19-47, figs. 1-57. [issn 0040-7496]. Published
30 November 1998.
The genus Heterothera Inoue, occurring widely in the Palaearctic and in the high mountains of
the Oriental region, is revised and twenty-three species are recognized. Nine species are de-
scribed as new: Heterothera hoenei Choi sp. n., H. yunnanensis Choi sp. n., H. triangulata Choi
sp. n., H. eclinosis Choi sp. n., H. stamineata Choi sp. n., H. obscurata Choi sp. n., H. distinc-
tata Choi sp. n., H. mussooriensis Choi sp. n. and H. kurenzovi Choi, Viidalepp & Vasjurin
sp. n., and five new combinations are suggested: Heterothera tephroptilus (Fletcher) comb, n.,
H. serrataria (Prout) comb, n., H. êtes (Prout) comb, n., H. comitahilis (Prout) comb. n. and
H. undulata (Warren) comb. n. A key and illustrations of adults and genitalia are given. The
monophyly of Heterothera and the species relationships are discussed. In addition, four poorly
known taxa of Thera sensu Prout, T. cyphoschema Prout (= atrinotata Joannis syn. n.), T. exan-
gulata Warren, and Pennithera distractata Sterneck comb, n., are redescribed and the taxono-
my of these species is briefly discussed.
Correspondence: Sei-Woong Choi, Department of Entomology, American Museum of Natural
History, Central Park West at 79th St., New York, NY 10024, USA. E-mail: choisw@amnh.org
Key words. - Systematics; Geometridae; Larentiinae; Heterothera; Palaearctic; Oriental.
The genus Heterothera Inoue consists of medium-
sized geometrid moths in the subfamily Larentiinae,
which are widely spread in the Palaearctic and Orien-
tal regions. The genus was erected by Inoue (1943)
based on the following characters: the absence of un-
cus, long anal tube, stout and flat saccus, weakly scle-
rotized costa and a well developed sacculus. The mo-
nophyly of the genus Heterothera s.s. was first defined
by Viidalepp (1980). He listed seven apomorphies for
the genus: the bifid saccus, the absence of an uncus,
the membranous ductus and corpus bursae without
signum, the simple sterigma, the distinct sacculus, the
presence of cornuti on the vesica, and the filiform
male antenna. Two synapomorphies out of seven, the
bifid saccus and the absence of uncus, were unique to
Heterothera s.s. However, the character 'absence of
uncus' was found to be incorrect.
Inoue (1982) proposed a new genus Viidaleppia for
the species of Asaphodes sensu Viidalepp (1980). Later,
he (Inoue 1986) described the characters of the genus
Viidaleppia as: doubly bipectinated male antenna,
strongly sclerotized costa and pointed apex of fore-
wing, sclerotized and plate-like sacculus, well devel-
oped apical projection of the sacculus, numerous spin-
ular cornuti, well developed sterigma, and a broad
ductus bursae.
However, the genus Heterothera s.s. was found to
be paraphyletic in relation to Viidaleppia and the lat-
ter was synonymized with Heterothera in a cladistic
analysis of the Cidariini sensu Herbulot (Choi 1997;
see 'Diagnosis and monophyly' for synapomorphies).
Since the works of Prout (1914, 1938, 1941), the
taxonomy of Thera s.l. has been considerably changed,
due to the discovery of many new species and a dif-
ferent analytical approach (e.g. Viidalepp 1980, Choi
1997). As a result of the cladistic analysis, the genus
Heterothera s.l. is characterized by several derived
characters and, based on these, many undescribed
species of the genus have been recognized, mainly
from southwestern China and northern India. The
purpose of the present study is to revise the species of
Heterothera. While the taxonomy of several poorly
known species of Thera s.l., such as T. exangulata, T.
cyphoschema, T. atrinotata and T. distractata, is uncer-
tain, another purpose is to redescribe these species in
order to understand their relationships better.
19
Tijdschrift voor Entomologie, volume i4i, 1998
Figs. 1-6. Adults of Heterothera. - 1 , H. postalbida; 2, H. tephroptilus; 3, H. mussooriensis; 4, H. un,
quadrifulta.
dulata; 5, H. hoenei; 6, H.
Materials and methods
The study is based on the material from the follow-
ing museums and private collections:
AMNH, American Museum of Natural History, New
York
BMNH, British Museum (Natural History), London
BNHM, Bulgarian Natural History Museum, Sofia
HNHM, Hungarian Natural History Museum,
Budapest
KY, private collection of Katsumi Yazaki, Tokyo
MF, private collection of M. Fibiger, Copenhagen
SNHM, Swedish Natural History Museum,
Stockholm
ZBi, Institute of Botany and Zoology, Tartu
ZFMK, Zoologisches Forschungsinstitut und
Museum Alexander Koenig, Bonn
ZMH, Zoological Museum, Helsinki
ZSM, Zoologische Staatssammlung München,
München.
Examination and dissection of the genitalia, in-
20
CHOI: Systematics ofHeterothera
eluding everting the vesica, follow Hardwick (1950),
while the terminology of the morphology including
the genitalia follows Forbes (1948) and Klots (1970).
Systematics
Heterotbera Inoue, 1943
Heterothera Inoue, 1943: 12. Type species: Cidaria postalbi-
d/zWileman, 1911 (original designation).
Viidaleppialnoue, 1982: 283. Type species: Cidaria quadri-
fiiltaVrout, 1938 (original designation).
Diagnosis and monophyly
Species of Heterothera are characterized by the scle-
rotized costa and the triangular sacculus of the male
genitalia, the well developed sterigma and the greatly
modified ductus bursae of the female genitalia. The
species of Heterothera have male antenna and a wing
pattern similar to the species of Pennithera Viidalepp,
Praethera Viiddalepp and Thera Stephens. However,
Heterothera can be distinguished from other taxa by
the triangular and sclerotized sacculus, several large
spinular cornuti, the well developed sterigma and the
greatly modified ductus bursae.
The monophyly of Heterothera s.l. has been de-
fined by Choi (1997), and seven synapomorphies
support the clade of Heterothera: (1) small process of
sacculus (or harpe), (2) long hairs on the cucullus, (3)
medially invaginated saccus, (4) scattered cornuti, (5)
semi-circular lamella antevaginalis, (6) relatively thick
ductus bursae, and (7) sclerites on the wall of the duc-
tus bursae.
Description
Antenna of male filiform {postalbida, yunnanensis,
sororcula, obscurata, eclinosis), bipectinate with short
pectinations (denti/asciata, distinctata, quadrifulta,
taigana, undulatd) or with long pectinations {incerta,
tephroptilns, consimilis, mussooriensis, firmata, serrarla,
serrataria, kurenzovi, hoenei). Frons smooth, covered
with blackish or dark ochreous and whitish scales.
Labial palp variable in length, often about twice as
long as eye diameter. Interantennal fillet dark brown-
ish in colour, often distinct by white scales. Legs
blackish or dark ochreous and whitish, with distinct
whitish tibial joints. Metathorax mediodorsally white,
with blackish tufts. Forewing ground colour varies;
basal part dark, occasionally white, basal line dentate,
slanted, occasionally smooth, vertical; dorsum be-
tween basal and antemedial lines with a black horizon-
tal streak {postalbida, sororcula, tephroptilus, êtes, mus-
sooriensis) or with a vertical blackish bar {yunnanensis,
taigana) or with a large dot {quadrifulta, consimilis);
antemedial line medially indented, occasionally not
indented, strongly waved; postmedial line costally
slanted, bent, often toothed, medially bulged; central
fascia variable in width, form constant throughout fas-
cia or thinner at middle and dorsum, discoidal dot
blackish, often indistinct by uniting with costal part of
antemedial line {êtes, firmata), dorsum distinct with
blackish scales {incerta, obscurata, dentifasciata, distinc-
tata, triangulata, quadrifulta, undulata, stamineata);
subterminal line sometimes present, blackish, scal-
loped {serraria, serrataria, kurenzovi); termen blackish
or dark ochreous. Hindwing ground colour whitish
{postalbida, yunnanensis, sororcula, incerta, dentifascia-
ta, distinctata, triangulata, consimilis, quadrifulta, ser-
raria, serrataria, kurenzovi, hoenei, stamineata, ecli-
nosis), yellowish white {êtes, firmata), greyish or
blackish {obscurata, tephroptilus, taigana, undulata, co-
mitabilis); discoidal dot usually small, often large;
postmedial line blackish, usually medially bulged; ter-
men sometimes tinged with blackish {incerta, triangu-
lata, consimilis, quadrifulta, serrataria, kurenzovi, un-
dulata) or with waving subterminal line {serraria) .
Male genitalia. - Uncus weakly sclerotized, tapered
at bottom or slender, length varies from moderate
{postalbida) to long {yunnanensis). Tegumen smaller
than the total length of vinculum and saccus, dome-
shaped or triangular. Saccus concave or medially in-
vaginated {postalbida, dentifasciata, yunnanensis, ob-
scurata, serraria, serrataria, kurenzovi, undulata,
eclinosis), broad and flat at bottom {incerta, tephrop-
tilus, distinctata, consimilis, taigana) or round {hoenei,
firmata). Juxta sclerotized, broad, occasionally juxta
and transtilla form a long tubular structure {postalbi-
da, sororcula); anellus lobe conspicuous, varies in
shape, from simple and digitiform {postalbida, soror-
cula, obscurata, distinctata, consimilis, firmata, taigana,
serrataria, kurenzovi, undulata, hoenei) to large and tri-
angular {yunnanensis, eclinosis) or small and nipple-
shaped {incerta, tephroptilus, dentifasciata, quadrifulta,
serraria), usually long hairs present at apex. Valva dis-
toventrally expanded, with long hairs; costa sclero-
tized, varies in width, distally wider {postalbida, soror-
cula, obscurata, tephroptilus, dentifasciata, distinctata,
consimilis, taigana, undulata, eclinosis), medially ex-
panded {yunnanensis, incerta, quadrifulta, hoenei,
mussooriensis) or medially and distally expanded {fir-
mata, serraria, serrataria, kurenzovi); sacculus distinct,
usually sclerotized, with a distal process {yunnanensis,
incerta, tephroptilus, dentifasciata, distinctata, consim-
ilis, firmata, quadrifulta, taigana, serraria, serrataria,
kurenzovi, hoenei, eclinosis) or two processes {postalbi-
da, sororcula, obscurata, undulata). Aedeagus sclero-
tized, slender, cylindrical or basally tapered to an
apex, distally scobinated, often large spinular process-
es around neck present {incerta, kurenzovi); vesica
tubular or small, sac-like, cornuti usually present,
variable in size, with from small to large spines.
Female genitalia. — Papillae anales weakly sclero-
21
Tijdschrift voor Entomologie, volume ui, 1998
i. ^ ^
10
14
Fig. 7-14. Adults of Heterothera. - 7, H. obscurata; 8, H. distinctata; 9, H. yunnanensis; 10, H. eclinosis; II, H. triangulata;
12, H. stamineata; 13, H. êtes; 14, H. kurenzovi.
22
CHOI: Sys tema tics of Heterotbera
tized. Segment 8 membranous or weakly sclerotized.
Anterior apophysis varies in length, from Vi to V$ of
posterior apophysis, often anterior apophysis missing
{taigana, serrarla, serrataria, kurenzovi). Ostium bur-
sae ventrally with lamella antevaginalis bar-shaped
(postalbida, yunnanensis, incerta, hoenei, stamineata,
eclinosis) or shell-shaped (triangulata); dorsally with
lamella postvaginalis forming large horn-like process-
es (taigana) or semi-round processes densely covered
with minute spines (serraria, serrataria, kurenzovi).
Ductus bursae varies in length, from very short {ser-
raria, serrataria, kurenzovi) to long, wall with sclerites
{postalbida, yunnanensis), sometimes largely expanded
from the anterior to the antrum {quadri/ulta). Corpus
bursae varies in size and shape between species, ovu-
lar, membranous (postalbida, incerta, denti/asciata) or
strongly sclerotized (serraria, serrataria, kurenzovi),
occasionally with appendix bursae (yunnanensis), of-
ten wall with minute scobinations; signum usually
absent, occasionally with one patch of scobinated
signa (taigana sounkeana).
Distribution
The species of Heterothera are widely distributed in
the Palaearctic and Oriental regions, but they are es-
pecially abundant in the Oriental region: there are
eight species in northern India and Nepal, six species
in southwestern China (Tibet), two species in Tai-
wan, five species in northeastern Asia, one species in
the western and eastern Palaearctic and one species in
the western Palaearctic only. Some species are quite
widely spread throughout the Palaearctic region. For
example, H. serraria is found from northern Europe
to eastern Siberia, H. taigana from central Siberia to
Kurile Islands and Japan, and H. postalbida from
western China to the Korean Peninsula and Japan.
On the other hand, many species are endemic to the
Oriental region (e.g. the two Taiwanese species H. in-
certa and sororcula).
Remarks
The bipectinated male antenna was used as a char-
acter for Viidaleppia (Inoue 1986). However, cladistic
analyses of the Cidariini (Choi 1997) and of Thera s A.
(Choi unpublished) indicate that the pectination of
male antenna is not a synapomorphy of Heterothera,
whereas the long pectination of the male antenna is a
good diagnostic character for some species of Hetero-
thera (e.g. incerta, tephroptilus, mussooriensis, hoenei).
The anellus lobes of the male genitalia in Hetero-
thera are quite helpful to diagnose taxa, especially
those from southwestern China. The lobes are usually
digitiform or rounded in the apical part. However,
two species, H. eclinosis and yunnanensis, possess great-
ly modified anellus lobes: they are apically sharply pro-
jected and medially triangular, expanded. Both species
occur in southwestern China. Nevertheless, the wing
pattern and the genitalia indicate that they are allied to
Heterothera and especially, the characters such as the
filiform male antenna and lamella postvaginalis of the
female genitalia show an affinity to H. postalbida.
The presence of spines around the neck of aedeagus
is recognized as a derived character for Thera (Vi-
idalepp 1980, Choi 1997). However, this feature is
also found in some species of Heterothera (e.g. incer-
ta, dentifasciata, hoenei, and yunnanensis) . Therefore,
it seems to me that the character is a symplesiomor-
phy for Thera s.l.
Checklist of Heterothera
Heterothera Inoue, 1943
H. postalbida (Wileman, 1911)
H. sororcula (Bastelberger 1909)
H. tephroptilus (Fletcher, 1961) comb. n.
H. mussooriensis sp. n.
H. undulata (Warren, 1888) comb. n.
H. hoenei sp. n.
H. incerta (Inoue, 1 986)
H. quadrifulta (Prout, 1938)
H. obscurata sp. n.
H. dentifasciata (Hampson, 1895)
H. distinctata sp. n.
H. yunnanensis sp. n.
H. eclinosis sp. n.
H. triangulata sp. n.
H. comitabilis (Prout, 1 923) comb. n.
H. stamineata sp. n.
H. consimilis (Warren, 1888)
H. êtes (Prout, 1926) comb. n.
H. taigana (Djakonov, 1926)
taigana taigana (Djakonov, 1926)
taigana sounkeana (Matsumura, 1 927)
taigana ishizukai (Inoue, 1955) syn. n.
H. serraria (Lienig, 1 846)
H. serrataria (Prout, 1914) comb. n.
H. kurenzovi Choi, Viidalepp & Vasjurin sp. n.
H. firmata (Hübner, 1822)
T'Aera Stephens, 1831
T. cyphoschema (Prout, 1926) incertae sedis
atrinotata Joannis, 1929 syn. n.
atrinotata reducta J oannis, 1929 syn. n.
T. exangulata (Warren 1909) incertae sedis
Pennithera Viidalepp, 1980
P. distractata (Sterneck, 1 928) comb. n.
23
Tijdschrift voor Entomologie, volume i4i, 1998
Figs. 15-20. Genital capsule in the male genitalia of Heterothera. -15, H. tephroptilus; 16, H. undulata; 17, H.
incerta; 18, H. hoenei; 19, H. obscurata; 20, H. dentifasciata.
Key to the species of Heterothera (males)
Two species, H. triangulata and H. stamineata, are
not included, because they are only known from fe-
males. However, based on the wing pattern elements,
H. triangulata is similar to H. dentifasciata and H. sta-
mineata is similar to H. comitabilis.
1. Antenna of male bipectinate 2
- Antenna of male filiform 16
2. Pectinations of male antenna long 3
- Pectinations of male antenna short 12
3. Dorsum of the forewing with a long black hori-
zontal streak 4
- Dorsum of the forewing without a black horizon-
tal streak 6
4. Postmedial line of the forewing costally ample
and round; costa of the valva nearly flat or distal-
ly expanded 5
- Postmedial line of the forewing costally slightly
invaginated; costa of the valva basally expanded..
H. mussooriensis
24
CHOI: Systematics of Heterothera
5. Costa of the valva nearly flat H. tephroptilns
- Costa of the valva distally widely expanded
H. êtes
6. Forewing ground colour blackish ....H. consimilis
- Forewing ground colour whitish or yellowish. ...7
7. Forewing ground colour whitish 8
- Forewing ground colour yellowish 10
8. Central fascia of the forewing medially reduced
or broken 9
- Central fascia of the forewing medially not re-
duced H. serrataria
9. Termen of the hindwing with a blackish waved
subterminal line; occurring in Europe, Central
and Eastern Siberia H. serrano.
- Termen of the hindwing without a blackish
waved subterminal line; occurring in the Russian
Far East, Korea or Japan H. kurenzovi
10. Forewing with a distinct blackish discoidal dot ...
11
- Forewing with a discoidal dot united to the costal
part of antemedial line H. firmata
1 1 . Central fascia of the forewing medially and dor-
sally reduced; costa of the valva medially slightly
expanded; ductus bursae relatively short, pleated
in general H. incerta
- Central fascia of the forewing nearly even in
width; costa of the valva medially triangular, ex-
panded; ductus bursae slim, membranous
H. hoenei
12. Postmedial line of the forewing costally strongly
slanting 13
- Postmedial line of the forewing costally relatively
less slanting 14
13. Central fascia of the forewing dorsally greatly re-
duced; interantennal fillet blackish
H. dentifasciata
- Central fascia of the forewing nearly constant in
width; interantennal fdlet distinct with white
scales H. distinctata
14. Central fascia nearly constant in width
H. taigana
- Central fascia reduced at dorsum 15
15. Antemedial line of the forewing medially sharply
indented; anellus lobe digitiform; ductus bursae
anterior to antrum greatly expanded
H. quadrifulta
- Antemedial line of the forewing medially not in-
dented; anellus lobe relatively thick; ductus bur-
sae anterior to antrum not expanded
H. undulata
16. Black horizontal streak or dot at dorsum between
basal and antemedial line of forewing present ..17
- No blackish scales at dorsum between basal and
antemedial line of forewing H. obscur ata
17. Presence of a black horizontal streak at the dor-
sum of forewine 18
- Presence of a black spot at the dorsum of
forewing 19
18. Postmedial line of the forewing costally slanting;
lamella antevaginalis reduced to small sclerotized
process; corpus bursae large H. sororcula
- Postmedial line of the forewing costally convex-
shaped; lamella antevaginalis well developed with
a pair of rod-shaped processes; corpus bursae rel-
atively smaller H. postalbida
19. Central fascia dorsally reduced; anellus lobe apical-
ly sharply projected and dentate, medially triangu-
larly projected; lamella postvaginalis with a pair of
large rod-shaped processes H. yunnanensis
- Central fascia relatively even in width; anellus
lobe apically sharply projected and bifid; lamella
postvaginalis simple H, eclinosis
Heterothera postalbida (Wileman)
(%• 1)
Cidaria postalbida Wileman, 1911: 325. Holotype â,
japan: Tokyo, Province]. Musashi, Honshu plains,
29.iv.1895, A.E. Wileman (bmnh) [examined].
Thera postalbida; Prout, 1941: 323.
Cidaria (Thera) postalbida problematica Bryk, 1948: 173.
Holotype $, korea: Juwool (Shuotsu), 28.vi.1935, (snhm)
[examined]. [Synonymized by Inoue, 1977: 268].
Heterothera postalbida; Inoue, 1943: 12.
Diagnosis
This species is distinguished by the greyish fore-
wing and whitish hindwing, blackish horizontal streak
at dorsum, smooth postmedial line, and the medially
reduced central fascia. The male genitalia show the
well developed complex of juxta and transtilla, digiti-
form anellus lobe with an expanded and angled apical
part, broad valva with two spinular sacculus processes
and apically densely scobinated aedeagus. In the fe-
male genitalia, the bar-like sclerotized lamella ante-
vaginalis and a bowl-shaped antrum are characteristic.
Redescription
Antenna of male filiform. Frons mixed with black-
ish brown and white, below projected. Labial palp
long, 1.5 times to twice of eye diameter. Wingspan
25-34 mm. Forewing ash grey; basal line grey, waved;
antemedial line smooth, dorsally greatly incurved;
postmedial line smooth, waved, costally and medially
bulged, subdorsally indented, dorsum projected; cen-
tral fascia reduced at bottom, discoidal dot long,
blackish; subterminal line whitish and undulating;
apical streak blackish; dorsum with long black hori-
zontal streak. Hindwing whitish, with grey discoidal
dot; fringe blackish. Thorax greyish, dorsum of
metathorax white with blackish tufts. Legs mixed
with blackish brown and white, tarsal joints distinct
and whitish.
25
Tijdschrift voor Entomologie, volume i4i, 1998
Male genitalia. - Uncus weakly sclerotized, moder-
ate, basally tapered. Tegumen dome-shaped, basally
incurved, with long, sclerotized anal tube. Saccus
broad, convex, sclerotized. Juxta and transtilla form a
tube-like complex, sclerotized, anellus lobe basally and
medially digitiform, apically expanded, triangular,
with long hairs. Valva distally broad, round with long
hairs; costa flat, sclerotized, sometimes distally ex-
panded; sacculus triangular, sclerotized, two distal
spinular projections. Aedeagus slender, distodorsally
densely scobinated; vesica tubular, with several minute
scobinated cornuti.
Female genitalia. — Papillae anales weakly sclero-
tized. Anterior apophysis Vi length of posterior apo-
physis. Antrum developed, deep, bowl-like, lamella
postvaginalis large bar-like sclerotized processes, la-
mella antevaginalis thin, round, sclerotized. Ductus
bursae medially twisted, with sclerotized walls. Cor-
pus bursae ovate, wall with very minute scobinations.
Distribution
Japan, Korea, Ussuri, SE China. Sterneck (1928)
reported one female from Guanxian, Prov. Sichuan
(Kwan-hsien).
Remarks
Inoue (1943) designated this species as the type of
Heterothera. He incorrectly described the uncus of the
male genitalia as absent.
Bryk (1948) proposed a new subspecies from Korea,
H. postalbida problematica. However, Inoue (1977)
synonymized the subspecies with the nominal species.
By checking the type specimens and the genitalia, I
confirm that subspecific division of the species is un-
warranted.
Four species, H. postalbida, sororcula, tephroptilus
and mussooriensis, are similar to each other in the
black horizontal streak along the dorsum of forewing,
wing ground colour and the genitalia. However, these
four species can be divided into two groups based on
the male antennal pectination and the geographic
range, (H. postalbida & sororcula) and (H. tephroptilus
& mussooriensis). Moreover, these species groups can
be distinguished by the shape of the dorsal part of sac-
culus, waved versus straight, and the processes of the
male genitalia, sharply pointed versus hooked.
Biology
The species is bivoltine (Wileman 1911) and in
Japan the larva is found only on Pinus densiflora (Sato
&Nakajima 1987).
Additional material examined. - korea: Kangwon Prov.
Mt. Kumkang-san, 26\ 26.V.1985. Dr. A. Vojnits & L.
Zombori; same locality, 2 9 , 17-19.ix.1980, Dr. L. Forro &
Gy. Topal; Pyongyang City, Pyongyang No. 1042. Mt. Ry-
oungak-san, 1 9, 13.X.1987, Korsos & Ronkay; S. Hwang-
hae Prov., Haeju, Mt. Suyong-san, No. 1050., 1$,
16.X.1987, Korsos & Ronkay (hnhm). - japan: Tokyo, 1 Î
10.V.1893, 15, 3.VÌ.1895, A.E. Wileman (bmnh), la,
l.v. 1953, P. Savolainen; Karuizawa, 1 ex., 24. vi. 1952. P.
Savolainen; Naganohara, 1 9, 9.x. 1969. J. Kaisila (zmh). -
china: Wenchow (Prov. Chekiang), 1$, 1913, C.T.
Bowring; Foochow (Prov. Fukien), la, April, 1885, Leech
(bmnh).
Heterothera sororcula (Bastelberger)
Thera sororcula Bastelberger, 1909a: 34. Holotype $, Tai-
wan: Arizan, Kagi-Distrikt 7-10000 Fuss, in Sencken-
berg Museum, Frankfurt [Colour transparency in bmnh,
examined] .
Dysstroma (Thera) sororcula; Bastelberger, 1909b: 172.
Thera sororcula; Prout, 1941: 323.
Heterothera sororcula; Inoue, 1992: 127.
Diagnosis
This species is very similar to H. postalbida in the
wing shape and the genitalia. However, it differs from
H. postalbida in the distinct central fascia, dark grey-
ish termen, less slanting antemedial line and angled
costal patt of postmedial line. In the female genitalia
the reduced lamella antevaginalis, strongly sclerotized
ductus bursae and large corpus bursae are distinguish-
ing features of H. sororcula.
Redescription
Antenna of male filiform. Frons covered with
btown and white scales. Labial palp long. Wingspan
27-29 mm. Forewing greyish, basal line blackish, den-
tate; area between basal and antemedial line tinged
with dark grey; antemedial line smooth, slanted; post-
medial line smooth, blackish, costally slanted, medial-
ly bulged, submedially indented, dotsum slightly
bulged; central fascia dark grey, with black discoidal
line, dorsally reduced; termen dark greyish, sometimes
whitish subterminal line appeared; apical streak black-
ish line; dorsum with black streak. Hindwing white
tinged with grey, with small discoidal dot; fringe
blackish. Thorax dark grey, dorsum of metathorax
white with black tufts. Legs mixed with brown and
white scales, tatsal joints white.
Male genitalia. - Uncus slendet, medially expand-
ed. Saccus broad, middle invaginated, anellus lobe
digitiform, apical patt not expanded, with long hairs.
Valva medio- and distoventrally membranous; costa
sclerotized, distally wider; sacculus triangular, sclero-
tized, one latge and one minute distal processes.
Aedeagus slender, distally scobinated; vesica tubular,
cornuti small spines, arranged into two layers.
Female genitalia. - Antrum bowl-like, lamella an-
tevaginalis bar-like, sclerotized processes, ventrally
thin and thread-like. Ductus bursae strongly sclero-
tized and twisted. Corpus bursae large, globular, wall
has minute scobinations.
26
CHOI: Systematics of Heterothera
Distribution
Taiwan.
Remarks
Bastelberger (1909a) noted that this species is sim-
ilar to H. consimilis Warren and suggested it might be
one subspecies of consimilis. However, he (1909b)
placed this species in Dysstroma based on the shape of
the male antennae. Prout (1941) noted its closeness
to H. postalbida.
Material examined. - Taiwan: Alishan (2200 m), Chiayi,
la 1°, IX.1964, Y.I.Chu; Id IS, 9-11. vii. 1964, H. In-
oue; Id, 19-22.vi.1943, M. Owada; Nan-Tou-sien, Id
1959, ex. H. Kezuka (bmnh); Tayuling (2600 m), Hualien
Hsien, Id, 26.iii.1987, Y. Shibata; AS 1?, l.iv.1984, A.
Kawabe; 2d 29, l.iv.1984, S. Sugi (bmnh); 19, 28-
29.viii.1983; ld, 2-4.V.1984; 1 d 19, 12-l4.iii.1985, H.
Yoshimoto (ky); Rantaizan, 7500 ft. 1 9 , 9.V.1909; Id,
10.V.1909; 19, 15.V.1909.A. E. Wileman (bmnh).
of posterior apophysis. Antrum sclerotized, simple,
relatively narrower. Ductus bursae broad, short, with
a sclerotized wall. Corpus bursae large, subspherical,
membranous.
Distribution
Nepal.
Remarks
The specimens from Gandaki (Nepal) are smaller
(wingspan 26-28 mm) than the type specimen
(wingspan 32 mm).
Additional material examined. — nepal: Paratypes ld,
Mustangbhot, 29°11' n. Br., 83°58' ö. L., Tagsa, 3500m,
12.viii.[19]55, Leg. F. Lobbichler; 1 9, same locality, 3800
m, 8.viii.[19]55 (bmnh); Mustangbhot, 29°11' n. Br.,
83°58' ö. L., Tagsa, 4300 m, 19, 24.viii.[19]55, F. Lob-
bichler (zsm); Gandaki above, Old Marpha 3450 m, 1 d
1 9 , 12.viii.1996, M. Fibiger (zmh).
Heterothera tephroptilus (Fletcher) comb. n.
(figs. 2, 15,27,46)
Them tephroptilus Fletcher, 1961: 170, figs. 10, 41, 50 &
51. Holotype d, nepal: Mustangbhot, 29°11' n. Br.,
83°58' Ö. L., Tagsa, 4300 m. 24.viii.[19]55, Leg. F. Lob-
bichler (zsm) [examined].
Diagnosis
This species is similar to H. postalbida in wing
shape, but differs in the long, bipectinate male anten-
na, blackish wing colour, the absence of the discoidal
dot, and medially deeply indented antemedial line of
forewing.
Redescription
Antenna of male bipectinate, with long pectinations.
Frons mixed with brownish and white. Labial palp
blackish, long. Wingspan 26-32 mm. Forewing black-
ish; basal line blackish; antemedial line costally greatly
slanted, dorsally waving; postmedial line costally slant-
ed, medially bulged; central fascia costally dark, dis-
coidal dot absent, medially reduced; dorsum with
black horizontal streak; apical streak black line. Hind-
wing blackish, with weak discoidal dot. Thorax dark
grey, dorsum of metathorax white with black tufts.
Male genitalia. - Uncus long, medially bent. Tegu-
men triangular. Saccus sclerotized, flat at bottom.
Juxta broad, sclerotized, anellus lobe short, round.
Valva mid- and distoventrally expanded; costally flat,
sclerotized; sacculus strongly sclerotized, triangular,
with dorsal border undulated, one sclerotized distal
spinular process. Aedeagus small, sclerotized surfaces,
having a bundle of small spines ventro- and distodor-
sally, small round vesica with several spinular cornuti,
arranged into a circle.
Female genitalia. - Anterior apophysis half length
Heterothera mussooriensis sp. n.
(figs. 3, 38, 39)
Type material. - Holotype cî, n. india: Mus-
soorie, 6000', Dehra Dun, U.P., 4.vii.l942 (amnh).
Paratype cT, same locality, 10.vii.1944 (amnh).
Diagnosis
This new species is very similar to H. tephroptilus in
the wing pattern and the male genitalia, but it differs
from tephroptilus in the slightly invaginated costal
part of the postmedial line of forewing, the discoidal
dot of hindwing, and the medially projected valval
costa of the male genitalia.
Description
Antenna of male bipectinate, with long pectina-
tions. Frons mixed with brown and ochreous scales.
Labial palp moderate, slightly longer than eye diame-
ter. Legs with yellowish tibial joints. Wingspan 27
mm. Forewing ground colour brownish; basal line
waved, medially bulged; antemedial line costally
blackish, discoidal dot united with costal part of line,
medially deeply indented, dorsally waved; postmedial
line costally slanted, medially bulged; central fascia
costally blackish, medially and dorsally reduced; dor-
sum with black horizontal streak; apical streak black
line; subterminal line very weak, waved. Hindwing
pale brownish; discoidal dot small, blackish.
Male genitalia. - Uncus long, slender, medially
bent. Tegumen small, dome-shaped. Juxta broad,
sclerotized; anellus lobe long, digitiform, subapically
expanded and fused with base of valva, apically long
hairs present. Saccus sclerotized, both edges slightly
projected. Ventral part of valva medially invaginated,
distally largely expanded; costa sclerotized, medially
27
Tijdschrift voor Entomologie, volume hi, 1998
projected; sacculus triangular, sclerotized, dorsal bor-
der waved, one distal spinular process. Aedeagus
cylindrical, distally scobinated; vesica small sac-like;
cornuti two bundles of small spines.
Female genitalia. — Unknown.
Distribution
N. India.
Etymology
The specific name refers to the type locality, Mus-
soorie, Northern India.
Heterothera undulata (Warren) comb. n.
(figs. 4, 16, 28)
Ypsitpetes undulata Warren, 1888: 326. Lectotype S , here
designated "n. india: Thundiani, 29.viii.[18]86." (bmnh).
Cidaria (Thera) undulata; P 'rout, 1938: 111.
Thera undulata; Prout, 1941: 323.
Diagnosis
H. undulata is identified by the blackish wings,
double vertical ante- and postmedial lines, and the
costally smooth and waving postmedial line of fore-
wing. The species is similar to H. hoenei, but it can be
distinguished from that by the blackish ground
colour of wings and medially less projected costa, an-
gled edges of saccus, and relatively thin sacculus of the
male genitalia.
Redescription
Antenna of male bipectinate, pectinations as short
as diameter of shaft. Frons mixed with brownish and
white. Labial palp as long as eye diameter. Wingspan
24-33 mm. Forewing dark grey; subbasai line present,
blackish, basal line blackish, slanted, costally project-
ed; antemedial line consisted of double vertical lines,
slanted, medially slightly indented; postmedial line
forming two vertical lines, smooth, medially bulged;
central fascia blackish, medially reduced, bottom dis-
tinct with blackish scales; termen blackish, subtermi-
nal line whitish, waved; apical streak black line.
Hindwing dark brownish, with blackish postmedial
line. Thorax with blackish tufts.
Male genitalia. - Uncus long and slender. Tegu-
men dome-shaped. Saccus strongly sclerotized, both
edges projected. Anellus lobe short, relatively thick in
width, with apically long hairs. Ventral part of valva
medially slightly invaginated; costa medially slightly
expanded; sacculus strongly sclerotized, triangular,
with distally two processes. Aedeagus basally tapered,
sclerotized, distally scobinated, vesica with three bun-
dles of cornuti.
Female genitalia. - Anterior apophysis Vi length of
posterior apophysis. Antrum broad; lamella antevagi-
nalis a pair of round projections, densely covered with
minute scobinations. Ductus bursae twisted, sclero-
tized. Corpus bursae ovate.
Distribution
India, Nepal.
Remarks
Hampson (1895) listed the species under the char-
acter "Antennae of male serrate and with fascicles of
cilia". Prout (1938) noted that H. undulata is similar
to Chloroclysta siterata Hufnagel or Hydriomena, but
differs in its antennal pectinations. Based on this, he
put the species under Thera.
Additional material examined. - N. india: Paralectotypes
2ô 1 9 , Thundiani, 6.ix.[18]86, (bmnh). - nepal: Ganda-
ki above, Old Marpha, 3450m, IcT 1Î, 12.viii.1996, M.
Fibiger (mf).
Heterothera hoenei sp. n.
(figs. 5, 18, 29,47)
Type material. - Holotype â , china: Lijiang (Li-
Kiang), Province Nord-Yunnan, 27.viii.1935, H.
Höne (zfmk). Paratypes 2â, same locality, 27.vii.
1935, I6.viii.1935, H. Höne; 19, Kangding (Ta-
tsien-Lou), Prov. Sichuan, 18.viii.1898, Chasseurs
indigènes (zfmk).
Diagnosis
H. hoenei is similar to H. undulata in the wing pat-
tern elements, but differs from the latter in the
strongly bipectinate male antenna, ochreous ground
colour of forewing, and whitish hindwing. In the
male genitalia, the species is characteristic by the me-
dially triangular process of costa, more or less round
saccus, and waving dorsal border of the sacculus and
its stellate process.
Description
Antenna of male bipectinate, with long pectina-
tions. Frons smooth, covered with white and blackish
scales. Labial palp ochreous, moderate about as long
as eye diameter, sometimes double of eye diameter.
Wingspan 24-29 mm. Forewing ground colour
ochreous; basal part whitish, basal line relatively
thicker, blackish, slanted, bordered with white thin
vertical line; area between basal and antemedial line
with blackish vertical bar-shaped band; antemedial
line blackish, medially indented; postmedial line
costally slightly curved, medially bulged; central fas-
cia whitish, sometimes tinged with blackish, discoidal
dot large, blackish, dorsally reduced; subterminal line
blackish, waved; apical streak blackish; fringe black-
ish. Hindwing whitish, with small blackish discoidal
28
CHOI: Systematic! of Heterotbera
dot; postmedial line blackish; fringe lined with black-
ish. Metathorax whitish, with black tufts.
Male genitalia. - Uncus moderate, medially bent.
Tegumen triangular. Saccus shallow, flat at bottom,
often midventraJly slightly invaginated. Juxta sclero-
tized, invaginated at bottom. Anellus lobe short, dig-
itiform. Valva distoventrally membranous, expanded;
costa sclerotized, medially triangular projection; sac-
culus relatively small, with disrally stellate process.
Aedeagus slender, sclerotized, medially bent; small
spinular cornuti present around neck of aedeagus.
Female genitalia. - Anterior apophysis Va length of
posterior apophysis. Lamella antevaginalis plate-like,
sclerotized processes, densely covered with minute
spines. Ductus bursae thin, long. Corpus bursae ovate.
Etymology
The specific name is given to honour H. Hone,
who dedicated his whole life to collecting and study-
ing Chinese Lepidoptera.
Distribution
S.W. China.
brown with whitish scalloped subterminal line; apical
streak black line. Hindwing whitish with black dis-
coidal dot; termen greyish with black fringe. Thorax
greyish, dorsum of metathorax white with black tufts.
Legs mixed with white and brownish black, tarsal
joints whitish.
Male genitalia. - Uncus slender, sclerotized, bent.
Tegumen dome-shaped. Saccus flat at bottom, sclero-
tized. Anellus lobe small, nipple-shaped. Ventral part
of valva medially expanded; costa medially expanded;
sacculus triangular, dorsal border slightly waved, with
one spinular process. Aedeagus cylindrical, slightly ta-
pered, distoventrally scobinated and distodorsally a
bundle of large spinular processes present; vesica
without cornutus.
Female genitalia. - Anterior apophysis Vi length of
posterior apophysis. Lamella antevaginalis a pair of
scobinated large lobes, ventrally semi-round plate
structure. Ductus bursae short, distally pleated. Cor-
pus bursae ovate, wall with minute scobinations.
Distribution
Taiwan.
Heterothera incerta (Inoue)
(figs. 17, 30)
Viidaleppia incerta Inoue, 1986: 234, figs 25D, 26. Holotype
S, Taiwan: Nengkaoshan, ca. 2800 m, Nantou Hsien,
20.V-2.VI. 1966, Ching-Shong Yu (bmnh) [examined].
Heterothera incerta; Choi, 1997: 311.
Diagnosis
H. incerta is identified by the bipectinate male an-
tenna, paler forewing, medially indented antemedial
line, the angled costal part of postmedial line, and the
distinct dorsum of central fascia and one patch of
spinular processes at the distal part of aedeagus. It is
similar to quadrifulta in the wing pattern, but incerta
is separated from the latter by having the blackish
scales at the bottom of the central fascia of forewing,
the waving dorsal border of sacculus, and the relative-
ly short and distally pleated ductus bursae.
Redescription
Antenna of male bipectinate, with long pectina-
tions, basal and distal parts ciliate. Frons mixed with
brownish black and white, below projected. Labial
palp long, nearly double of eye diameter. Wingspan
26-27 mm. Forewing greyish or dark greyish; sub-
basal, basal lines blackish; area between basal and an-
temedial lines with reddish brown band; antemedial
line costally slanted, medially indented, below slight-
ly dentate; postmedial line costally slightly angled,
slanted, medially bulged, dorsally waved; central fas-
cia dark greyish, dorsally reduced, blackish scales at
bottom, discoidal dot blackish; termen reddish
Remarks
Inoue (1986) noted that this species was similar to
Dysstroma citratum (Denis & Schiffermüller) [incor-
rect authorship, Linnaeus] on the basis of wing shape,
colour and maculation. He also noted that this species
was similar to quadrifulta in the male genitalia.
Material examined. - Taiwan: Mt. Yu Shan (3528 m),
Paiyunshanchuang, Kaohsiung Hsien, 1 S 1 $ , 2-3-viii. 1985,
M. Nishizawa (ky); Hohuanshan, 3100m, Nantou Hsien,
1 6 1 $ , 5.viii.l987, A. Kawabe, Coll. Inoue; Nantou Hsien,
Formosa, 1 9 , 1968, ex. H. Kezula, Coll. Inoue (bmnh).
Heterothera quadrifulta (Prout)
(fig. 6)
Cidaria (Thera) quadri '\fi dtaV 'rout, 1938: 114. Lectotype â,
here designated "japan: Shinano, 2.viii. 1911, coll[ection]
Wileman" (bmnh).
Asaphodes quadrifulta; Viidalepp, 1980: 64.
Viidaleppia quadrifidta; Inoue, 1982: 283.
Heterothera quadrifulta; Choi, 1997: 311.
Diagnosis
The wings of quadrifulta have whitish ground colour
with shades of black, a waving antemedial line at the
costal part, blackish discoidal dot, white scalloped sub-
terminal line and basally blackish tinged hindwing.
This species is similar to H. incerta, but differs in the
whitish ground colour of forewing and short pectina-
tions of male antenna. Distinguished from H. incerta
by the middorsal expansion of the costa in the male
genitalia and the distally largely expanded ductus bur-
sae of the female genitalia.
29
Tijdschrift voor Entomologie, volume i4i, 1998
Figs. 21-26. Genital capsule in the male genitalia of Heterothera. — 21, H. distinctata; 22, H. yunnanensis; 23, H. eclinosis;
24, H. firmata; 25, H. serrataria; 26. H. kurenzovi.
Redescription
Antenna of male bipectinate, with short pectinations,
about same width as the shaft. Wingspan 31 mm.
Forewing ground colour whitish; basal part dark grey,
basal line blackish, slightly dentate; area between
basal and antemedial lines with dark grey band; ante-
medial line costally extended inward, medially in-
dented, dorsally slightly extended inward; postmedial
line costally slanted, medially projected, below waved;
central fascia greyish, with black discoidal dot, medi-
ally reduced, bottom distinct with blackish scales;
subterminal line costally blackish, medially whitish,
undulating; termen dark grey. Hindwing whitish, ter-
men blackish, with black discoidal dot. Thorax mixed
with brownish and white scales with tufts.
Male genitalia. - Uncus long, sclerotized, basally
tapered. Tegumen dome-shaped. Anellus lobe small,
nipple-shaped. Ventral part of valva medially slightly
invaginated; costa sclerotized, middorsally expanded;
sacculus triangular, dorsal border nearly straight, with
30
CHOI: Systematic! of Heterothera
one large distal process. Aedeagus disrally scobinated;
vesica with two bundles of cornuti.
Female genitalia. - Anterior apophysis Vi length of
posterior apophysis. Lamella antevaginalis tongue-
shaped with a pair of round patch and densely scobi-
nated. Ductus bursae long, greatly expanded at distal
part, wall with rows of spines and sclerites. Corpus
bursae membranous, ovate, signum absent.
Distribution
Korea, Japan.
Remarks
Prout (1938) described the species from two males
from Shinano, Japan. He noted that the species is sim-
ilar to Pennithera comis Butler and Heterothera taigana
sounkeana Matsumura, but differs from P. comis in the
less distinct dorsal dot between basal and antemedial
line of forewing. Inoue (1982) designated this species
as type species for Viidaleppia.
Additional material examined. - japan: Ontake, Gifu-
ken. 1 S , 12-15.viii.1936, coll. Wileman; Kobushi, Saitama,
ld, 4.VÜL1953, T. Haruta (bmnh); Mt. Norikuradake,
2600m, Azumi-mura, Nagano, Id, 6.viii.l988, Y. Kishida;
same locality, 19, 17.viii.1988, K. Yazaki; Odarumi-toge,
Yamanashi, Id 1 9, 29.vii.1990, K. Yazaki (ky). - korea:
Prov. Ryang-gang, Chann-Pay Plateau, Sam-zi-yan, 1700m,
No. 285, Id, 24.vii.1975, J. Papp & A. Vojnits (hnhm).
Male genitalia. - Uncus long, sclerotized, medially
bent. Tegumen bell-shaped, sclerotized. Saccus scle-
rotized, medially broadly invaginated. Juxta sclero-
tized, anellus lobe long, digitiform with hairs on top.
Costa of valva strongly sclerotized, distally wider; sac-
culus triangular, sclerotized, distally two processes
with one large and one small. Aedeagus short, cylin-
drical, distally tapered and scobinated; vesica small,
with several large and small spinular cornuti.
Female genitalia. - Unknown.
Etymology
The specific name refers to the indistinct pattern
elements of forewing.
Distribution
Nepal.
Heterothera denti/asciata (Hampson)
(figs. 20, 32)
Larentia denti/asciata Hampson, 1895: 379. Lectotype d,
here designated "Pakistan: Murree, 1884, Harford
coll[ection]." (bmnh).
Cidaria {Therd) denti/ascia; Prout, 1914: 219. [Incorrect
spelling].
Thera denti/asciata; Prout, 1941: 324.
Viidaleppia dentifasciata; Yazaki, 1994.
Heterothera dentifasciata; Choi, 1 997: 311.
Heterothera obscurata sp. n.
(figs. 7, 19,31)
Type material. - Holotype cT, w. nepal: Karnali,
Mugu District]., Rara lake 2,990 m, 25- 26.ix.1981,
M. Owada, Coll. K. Yazaki (zfmk).
Diagnosis
This species is quite similar to incerta in wing pattern
elements but can be separated from it by the filiform
male antenna without pectinations, its greyish wings
and by the male genitalia which have a relatively long
anellus lobe, a distally expanded costa, a relatively thin
sacculus, and scattered spinular cornuti of the vesica.
Description
Antenna of male filiform. Frons covered with
white and blackish scales. Labial palp long, about 1.5
times of eye diameter. Metathorax has blackish tufts.
Wingspan 27 mm in male. Forewing ground colour
greyish; basal line dentate; antemedial line medially
sharply indented; postmedial line costally slanted,
waved, medially bulged; central fascia greyish, with
black discoidal dot, blackish scales present at bottom,
medially reduced; termen brownish, with weak black-
ish subterminal line. Hindwing greyish, with blackish
postmedial line.
Diagnosis
H. dentifasciata is easily identified by the blackish
and greatly slanting costal part of postmedial line and
large blackish discoidal dot of forewing. The male
genitalia of dentifasciata are similar to those of H. ob-
scurata, but the processes of sacculus and the cornuti
of the vesica are different.
Redescription
Antenna of male bipectinate, with short pectina-
tion as long as shaft. Frons mixed with brownish
black and white scales. Labial palp long about twice
of eye diameter. Wingspan 29-34 mm. Forewing red-
dish brown or greyish; basal line blackish; antemedial
line weakly slanted, medially indented; postmedial
line costally strongly slanted, medially bulged; central
fascia blackish, medially broader and dorsally re-
duced, with black discoidal dot; subterminal line
blackish and undulating; apical streak blackish line.
Hindwing whitish, basal slightly blackish, with black
discoidal dot and postmedial line, fringe black. Meta-
thorax middorsally white with black tufts.
Male genitalia. - Uncus long, slender, sclerotized,
basally tapered. Tegumen dome-shaped. Saccus scle-
rotized, medially concave- shaped. Anellus lobe small,
digitiform. Ventral part of valva distally wider, with
long hairs; costa sclerotized, distally expanded; saccu-
31
Tijdschrift voor Entomologie, volume i4i, 1998
li
34
35
32
CHOI: Systematics of Heterothera
lus triangular, with a distal spinular process. Aedeagus
sclerotized, basally tapered, distally scobinated; vesica
with two bundles of cornuti.
Female genitalia. - Anterior apophysis half length
of posterior apophysis. Antrum broad, strongly scle-
rotized. Ductus bursae twisted, relatively short, thick.
Corpus bursae small, ovate.
Distribution
India, Pakistan, Nepal.
Remarks
Hampson (1895) listed the species with Pennithera
comis under the character "Antennae of the male with
two pairs of short cilia-bearing processes from each
joint". Prout (1938) noted that the species is similar to
consimilis in the colour of central fascia and it is also
similar to Thera exangulata^f] arren, but differs from T.
exangulata by the less waved basal line of forewing, pale
hindwing and the angled postmedial line of hindwing.
Additional material examined. - nepal: Central N.
Ganesh Himal. Nesim 2200m, 26, 23.x. 1995, M. Fibiger;
Central N. Ganesh Himal. Godlang 2560 m, la 2 $ ,
13.x. 1995, M. Fibiger; Sagarmatha N.P., 27°49'N 86°44'E,
Kyang Juma, 3600 m, 1Î, 19.V.1996, A. Albrecht, O.
Bistrom, K. Mikkola & A. Wikberg; Sagarmatha N.P.,
27°48'N 86°44'E, Syanboche, 3800 m, 2Î, 22.V.1996, A.
Albrecht, O. Bistrom, K. Mikkola & A. Wikberg (zmh); W.
Nepal, Karnali, Mugu Dist. Rara Lake, 2990 m, là, 25-
26.ix.1981, M. Owada (ky); Bagmati Rasuwa, Yuli Karka,
3375m, 19, 12-13.V.1993, T. Haruta(KY).
Heterothera distinctata sp. n.
(figs. 8, 21,33)
Type material. - Holotype 6, china: Lijiang (Li-
Kiang), Province Nord-Yunnan, 21.iv.1935, H. Hone
(zfmk).
Diagnosis
The new species is identified by the short pectina-
tions of the male antenna, whitish interantennal fillet,
ochreous forewing, whitish hindwing, and the strong-
ly slanted costal part of the postmedial line of fore-
wing. This new species is very similar to denti/asciata in
the wing shape and the male genitalia, but it can be dis-
tinguished from dentifasciata by the whitish interan-
tennal fillet and the even width of central fascia, which
is not reduced at the middle and dorsum of fascia.
Description
Antenna of male bipectinate, with short pectina-
tions. Interantennal fillet and frons white. Labial palp
moderate. Legs dark ochreous, with whitish tibial
joints. Metathorax with whitish hairs on middorsum,
with black tufts. Wïngspan 29 mm in male. Forewing
ground colour yellow ochreous; basal line blackish,
nearly vertical; area between basal and antemedial line
tinged with blackish scales; antemedial line blackish,
medially indented; postmedial line costally slanted,
medially bulged; central fascia ochreous, discoidal dot
one large blackish, dorsally distinct with blackish
scales; subterminal line blackish and undulating; apical
streak blackish line. Hindwing whitish, with black dis-
coidal dot; postmedial line blackish, medially bulged.
Male genitalia. - Uncus sclerotized, medially bent,
moderate in length, as long as height of tegumen.
Tegumen triangular. Saccus sclerotized, broad and
flat at bottom. Juxta sclerotized. Anellus lobe small,
digitiform, with hairs on top. Costa sclerotized, dis-
tally largely expanded; sacculus triangular, sclero-
tized, distally with a hooked process. Aedeagus distal-
ly tapeted, sclerotized; vesica small, sac-like, cornuti
mixed with small and medium spinular processes, lo-
cated around neck and middle of vesica.
Female genitalia. - Unknown.
Etymology
The specific name refers to the diagnostic wing
pattern elements of the forewing.
Distribution
S.W. China.
Heterothera yunnanensis sp. n.
(figs. 9, 22, 34, 48)
Type material. - Holotype S , china: Lijiang (Li-
Kiang), Province Nord-Yunnan. 27.vi.1935, H. Hone
(zfmk); Paratype 9 , china: A-tun-tse (Nord-Yunnan),
Talsohle ca. 3000m, 4.vii.l937, H. Hone (zfmk).
Diagnosis
H. yunnanensis is distinguished by the blackish and
greatly reduced central fascia at the middle and dorsal
parts and the black horizontal streak along the dorsum
of basal part and central fascia. The anellus lobes of the
male genitalia, apically sharp and pointed, medially tri-
angular and expanded, are characteristic. This new
species is quite similar to H. dentifasciata in wing shape,
Figs. 27-37. Aedeagus with everted vesica of Heterothera. - 27, H. tephroptilus; 28, H. undulata; 29, H. hoenei; 30, H. incer-
ta; 31, H. obscurata; 32, H. dentifasciata; 33, H. distinctata; 34, H. yunnanensis; 35, H. eclinosis; 36. H, serrataria; 37, H.
kurenzovi.
33
Tijdschrift voor Entomologie, volume i4i, 1998
Figs. 38-45.
Male genitalia of Heterothera
and Thera s.l.; 38, 40, 42, 44,
genital capsule; 39, 41, 43, 45,
aedeagus. - 38, 39, H. mussoor-
iensis; 40, 41, 7! cyphoschema;
42, 43, J", exangukta; 44, 45,
Pennithera distractata.
Scale bars: 1 mm.
34
CHOI: Systematics of Heterothera
but it can be distinguished from H. denti/asciata by the
filiform male antenna, black streak at the dorsum of
forewing, the anellus lobe and the cornuti of the vesica.
The lamella antevaginalis of the female genitalia, with a
rod-shaped process, is similar to H. postalbida.
Description
Antenna of male filiform. Frons mixed with white
and blackish brown scales. Labial palp moderate in
length. Legs mixed with white and blackish brown.
Metathorax blackish, with black tufts. Wingspan 32
mm in male, 36 mm in female. Forewing ground
colour greyish; basal line blackish, consisting of dou-
ble vertical lines, dentate; antemedial line medially
slightly indented, slanting from costa to dorsum;
postmedial line costally smooth, slanted, medially
bulged, dorsally waving; central fascia blackish, medi-
ally and dorsally greatly reduced, dorsally blackish;
border of costal part of postmedial line lined with
whitish; subterminal line blackish and undulating;
termen greyish; apical streak black line; dorsum be-
tween basal and antemedial lines with black bar.
Hindwing whitish; postmedial line blackish, double
vertical lines; fringe dotted with black.
Male genitalia. - Uncus slender, long, about 1.5
times the height of tegumen. Tegumen bell- shaped.
Saccus medially invaginated. Juxta round at bottom,
sclerotized. Anellus lobe medially triangular, sclero-
tized, apically sharp, slightly bifurcated. Costa sclero-
tized, medially wider; sacculus triangular, sclerotized,
distally spine-like process. Aedeagus slender, sclero-
tized; cornuti comprised of three bundles of spinular
processes at ventro- and dorsobasal part and middle
part of vesica.
Female genitalia. - Papillae anales weakly sclero-
tized. 8th segment weakly sclerotized. Lamella ante-
vaginalis laterally a pair of large spinular spines, ven-
trally a pair of ring-like sclerotized processes with
densely covered small spines. Ductus bursae short,
with sclerotized wall. Corpus bursae large with one
large appendix bursae.
Etymology
The specific name refers to the type locality, Yunnan.
Distribution
S.W. China.
Heterothera eclinosis sp. n.
(figs. 10, 23, 35, 49)
Type material. - Holotype S , china: Taibaishan
(Tapaishan im Tsinling), Sued-Shensi, 30.vi.1935, L.
(1700m), H. Höne (zfmk); Paratype 9 , same locali-
ty, 16.V.1936, H. Höne (zfmk).
Diagnosis
H. eclinosis is characterized by the slanted and dor-
sally blackish central fascia of forewing. It is similar to
H. yimnanensis in the wing pattern and the male gen-
italia, but it can be distinguished by its smaller size,
ochreous ground colour of wings and relatively con-
stant width of central fascia and the medially not ex-
panded anellus lobe of the male genitalia and the gen-
eral structure of the female genitalia.
Description
Antenna of male filiform. Frons with white and
blackish scales. Labial palp long, about 1.5 times the
eye diameter. Legs dorsally blackish, with whitish tib-
ial joints. Thorax dorsally white, metathorax with
blackish tufts. Wingspan 27 mm in male, 30 mm in
female. Forewing ground colour ochreous; basal area
dark ochreous, basal line waving; antemedial line
greatly slanted, medially indented; postmedial line
medially bulged, slightly waving; central fascia dark
ochreous, dorsally slightly reduced; termen blackish,
subterminal line whitish, undulating; apical streak
blackish line; dorsum with black dot. Hindwing
white, with small discoidal dot; postmedial line light
blackish, medially bulged; fringe black.
Male genitalia. - Uncus strongly sclerotized, long,
about twice of height of tegumen. Tegumen triangu-
lar. Saccus sclerotized, medially invaginated. Juxta
sclerotized. Anellus lobe sclerotized, long, rod-shaped,
medially elbowed, apically bifurcated. Costa sclero-
tized, distally wider; sacculus triangular, sclerotized,
with a distal spinular process. Aedeagus slender, scle-
rotized; cornuti small spinular process, present on
basal and middle parts of vesica.
Female genitalia.- Anterior apophysis Vi length of
posterior apophysis. Lamella antevaginalis a pair of
triangular plates which are densely covered with small
spines. Antrum thin, sclerotized. Ductus bursae rela-
tively short, sclerotized. Corpus bursae with ovate ap-
pendix bursae, without signum, wall has minute sco-
binations.
Etymology
The specific name refers to the sloping central fas-
cia of the forewing.
Distribution
S.W. China.
Additional material examined. - china: Taibaishan
(Tapaishan im Tsinling), Sued- Shensi, lo* , 2.VÜ.1935, 1 9
23.vi.1935, H. Höne. (bmnh).
35
Tijdschrift voor Entomologie, volume i4i, 1998
Heterothera triangulata sp. n.
(figs. 11,50)
Type material. - Holotype 9 , china: Lijiang
(Likiang), Province Nord-Yunnan, 25.ix.1935, H.
Höne (zfmk).
Diagnosis
This species is very similar to H. dentifasciata in the
central fascia. However, it can be distinguished from
the latter by its smaller wings, less indented anteme-
dial line, and the lamella antevaginalis and ductus
bursae of the female genitalia.
Description
Antenna of female filiform. Frons mixed with
white and blackish scales, with blackish projecting
hairs below. Labial palp long, double the eye diame-
ter. Legs mixed with white and blackish scales, and
whitish tibial joints. Metathorax whitish at middor-
sum, with blackish tufts. Wingspan 27 mm in female.
Forewing ground colour ochreous; basal part whitish,
basal line relatively thicker, blackish; antemedial line
smooth, medially slightly indented; postmedial line
costally strongly slanted, medially bulged; central fas-
cia tinged with blackish scales, with large discoidal
dot, medially reduced, dorsally distinct and blackish;
subterminal line weak, whitish vertical line; apical
streak blackish line. Hindwing whitish, with small
black discoidal dot; blackish postmedial line, medial-
ly bulged; termen blackish.
Male genitalia. - Unknown.
Female genitalia. - 8th segment long, with weakly
sclerotized surface. Lamella antevaginalis shell-like pro-
cesses. Antrum thin, sclerotized. Ductus bursae rela-
tively thin, long. Corpus bursae ovate, without signum.
Etymology
The specific name refers to the triangular central
fascia of forewing.
Distribution
S.W. China.
Heterothera comitabilis (Prout) comb. n.
Larentia comis Hampson, 1895: 379 (in part), nee Butler.
Them comitabilis Prout, 1923: 198. Holotype cT, n. india:
Sikkim, Jongri, 13000 ft., 1887, ex. coll. Elwes (bmnh)
[examined] .
Diagnosis
The waved costal part of postmedial line, the rela-
tively thicker and the blackish subterminal line of
forewing and blackish hindwing are characteristic fea-
tures of comitabilis. The species is similar to H. sta-
mineata, but differs in the costally toothed postmedi-
al line of forewing and blackish ground colour of
hindwing.
Redescription
Antenna of male bipectinate with short pectina-
tions. Frons covered with dark ochreous and white
scales. Labial palp long, double of eye diameter. Inter-
antennal fillet whitish. Wingspan 30 mm. Forewing
dark ochreous; basal part blackish, basal line distinct
with whitish; area between basal and antemedial line
with blackish bar; antemedial line medially indented;
postmedial line costally dentate, slightly tinged with
white, medially bulged; central fascia blackish, with
black discoidal dot; subterminal line thick, blackish,
undulating. Hindwing pale blackish; discoidal dot
blackish; postmedial line paler; fringe black.
Male genitalia and female genitalia not examined
(see remarks below).
Distribution
N. India.
Remarks
Prout (1923: 198) noted that the species is similar
to Pennitbera comis Butler in the antennal structure,
but differs in the short branches of antenna. He also
noted that the species is similar to Electrophaes coryla-
ta in the darkish coloration.
The genitalia of comitabilis are not examined be-
cause the types listed were lacking their abdomens.
The placement of the species in Heterothera is based
on the shape of wing pattern elements which is simi-
lar to H. stamineata.
Additional material examined. - Paratype 5 , N. india.
Sikkim, Tonglo, July 1886, Collectio[n] H.J. Elwes. (bmnh).
Heterothera stamineata sp. n.
(figs. 12, 51)
Type material. - Holotype 9 , china: Lijiang (Li-
kiang), Province Nord-Yunnan, 4.vii. 1935, H. Höne
(zfmk); Paratype 9 , same locality, O, 17.vi.1935, H.
Höne (zfmk).
Diagnosis
The new species is similar to comitabilis in the
shape of the wings, but differs in the whitish ground
colour of the hindwing.
Description
Antenna of female filiform. Frons covered with
white and blackish scales. Labial palp length twice the
eye diameter. Metathorax with blackish tufts. Wing-
span 30-31 mm in female. Forewing ground colour
brownish; basal part dark brownish, basal line den-
36
CHOI: Systematics of Heterothera
tate; antemedial line bordered with whitish, costally
slanted, medially and dorsally nearly vertical; post-
medial line costally bordered with whitish line, medi-
ally bulged; central fascia brownish, discoidal dot
black, bottom blackish; subterminal line blackish,
undulating; fringe dotted with blackish. Hindwing
whitish, with black discoidal dot; postmedial line
blackish; fringe lined with black.
Male genitalia. — Unknown.
Female genitalia. — 8th segment weakly sclerotized.
Anterior apophysis Va length of posterior apophysis.
Lamella antevaginalis a plate-like process, densely cov-
ered with small spinular spines, ventrally with semi-
circular sclerotized plate. Ductus bursae relatively
short, thin. Corpus bursae ovate, without signum.
Etymology
The specific name refers to the shape of the thread-
like central fascia.
Distribution
S.W. China.
Heterothera consimilis (Warren)
(fig. 52)
Thera consimilis Warren, 1888: 326. Holotype 8 , n. india:
Thundiani, 9.x. [18] 86, Genitalia slide No. 3986. (bmnh)
[examined]
Larentia consimilis; Hampson 1895: 380.
Heterothera consimilis; Choi 1997: 311.
Diagnosis
H. consimilis is easily identified by the wavy central
fascia and dentate basal line of the forewing. In the fe-
male genitalia the thick and sclerotized ductus bursae is
characteristic. This species is similar to H. êtes in wing
pattern, but differs in the blackish ground colour of
Redescription
Antenna of male bipectinate, with long pectina-
tions, tip and base filiform. Frons mixed with brown-
ish and white scales. Labial palp long, twice diameter
of eye. Thorax mixed with brownish and white scales,
with tufts. Wingspan 24-29 mm. Forewing ground
colour dark greyish or blackish; veins of basal part on
blackish streak; subbasai line blackish, round; basal
line black, dentate; area between basal and antemedi-
al line with weak blackish band; antemedial line
costally slanted, medially and dorsally waved; post-
medial line costally straight, often bent, slanting, me-
dially bulged, below waving, dorsally bulged; central
fascia dark grey, with black discoidal line, sometimes
medially and dorsally reduced; dorsum with large
reddish dot; termen dark greyish with white undulat-
ing subterminal line; apical streak a black line. Hind-
wing whitish, termen dark greyish, with small black
discoidal dot.
Male genitalia. - Uncus long, sclerotized, and
slightly bent in middle. Tegumen triangular. Saccus
flat at bottom and sclerotized. Anellus lobe small, dig-
itiform. Ventral part of valva distally slightly expand-
ed; costa sclerotized, medially very wider; sacculus tri-
angular, dorsal border waved, with one distal large
sclerotized spine. Aedeagus distally scobinated, with
round vesica and cornuti in two bundles.
Female genitalia.- Anterior apophysis Vi length of
posterior apophysis. Antrum simple, unmodified.
Ductus bursae thick, wall sclerotized. Corpus bursae
ovate, small.
Distribution
India, Nepal, Afghanistan.
Remarks
Warren (1888) described the species based on five
males and one female from Thundiani and noted that
the species is closely related to Thera juniperata. Prout
(1938, 1941) wrote that this species is similar to T.
cupressata Herrich-Schäffer in the wing colour and he
also pointed out the colour and size variation among
the specimens of Thundiani, Kashmir and Simla.
Additional material examined. - india: Paratype 1 9 ,
Thundiani, 2.v. [ 1 8] 87, Genitalia slide No. 3987; Simla, 7000
ft., 3d 29 A.E. Jones; Narkundah, 19, April. 1888, H.
McArthur coll.; Goorais Valley, Id, Junel887, J.H. Leech;
Kashmir, Gulmorg, 3d, 10.vi.[19]31, 12.vi.[19]3L 16. vi.
[19]31, Fletcher coll. (bmnh); West Bengal, Darjeeling Dist.
Tonglu, 3040 m, 29, 6.XÌ.1981, M. Owada (ky); N-Indien,
Uttar Pradesh, Kumson Himalaya, Dist. Nainital, Bhimtal,
1500m, 19, 15-30.V.1990, A. Hauenstein (zfmk). - nepal:
Mustangbhot, 29° 11' n. Br., 83°58' ö. L, Tagsa, 3500 m.
9.vii.[19]55, F. Lobbichler (bmnh); Gandaki above, Old
Marpha. 3450m, Id, 12.viii.1996, M. Fibiger; 15 km SW,
Kathmandu Hattiban pine for., 1500m, 19, 2.iii.l995, K.
Mikkola & A. Wikberg; Godavari, 27°40'N 85°25'E, 15 Km
SE Kathmandu, 1500 m, 19, 25.V.1996, A. Albrecht, O.
Bistrom, K. Mikkola & A. Wikberg (zmh); Kathmandu, Go-
davari 1600m, Id, 18.vii.1990, K. Yazaki (ky). - Afghan-
istan: Afghanistan, 1 9 , 84-41, Fletcher coll. (bmnh).
Heterothera êtes (Prout) comb. n.
(figs. 13, 53)
Thera êtes Prout, 1926: 251. Holotype 8, n. india: Assam
5000ft. Shillong, November] 1924, Fletcher leg. (bmnh)
[examined]
Diagnosis
The species is very similar to H. consimilis in the
wings and genitalia, but differs in the greyish wing
ground colour, the process of sacculus and the three
bundles of cornuti of the vesica.
37
Tijdschrift voor Entomologie, volume i4i, 1998
46
47
*S*i
50
51
Figs. 45-5 1 . Female genitalia of Heterotbera. - 46, H. tephroptilus; Al, H. hoenei; 48, H. yunnanensis; 49, H. eclinosìs; 50, H.
triangulata; 51, H. stamineata.
Redescription
Antenna of male bipectinate, with long pectinations.
Frons mixed with yellowish and yellowish brown
scales, smooth. Labial palp long, about 1.5 times of eye
diameter. Legs yellowish and yellowish brown. Meta-
thorax with blackish tufts. Wingspan 26 mm. Fore-
wing ground colour greyish; basal line blackish, den-
tate; antemedial line costally blackish, largely waved
at middle and dorsum, medially indented; postmedi-
al line blackish, medially and dorsally bulged; central
fascia waved, discoidal dot long and distinct by unit-
ing with costal part of antemedial line; subterminal
line indistinct, pale whitish; apical streak light black-
ish; dorsum with black horizontal streak. Hindwing
yellowish white; blackish postmedial line.
Male genitalia. - Uncus sclerotized, long about
twice of tegumen height. Tegumen dome-shaped.
Saccus broad, sclerotized, medially slightly invaginat-
ed. Juxta broad, sclerotized; anellus lobe long, digiti-
form, apically expanded with long hairs. Ventral part
of valva medially invaginated, distally wide with a
patch of hairs; costa sclerotized, medially much wider;
sacculus triangular, dorsal border waved, with one dis-
tal process. Aedeagus cylindrical, sclerotized, medially
slightly bent, distally densely scobinated; vesica small
sac-like, cornuti spinular processes in three patches.
38
CHOI: Systematics of Heterothera
Female genitalia. - Papillae anales weakly sclero-
rized. Anterior apophysis 1/5 length of posterior apo-
physis. Antrum simple. Ductus bursae moderate, wall
with sclerotized process. Corpus bursae ovate, signum
absent.
Distribution
N. India.
Remarks
Prout (1926) noted that the species is similar to Pen-
nithera comis Butler and H. consimilis Warren. Howev-
er, he distinguished êtes based on its smaller size, red-
dish and weakly marked wing pattern elements and
slender male abdomen. He (1941) noted that the
species is somewhat similar to Them cyphoschema Prout
or T. atrinotata Joannis in the wing pattern, but it can
be distinguished by the male antennal pectination.
Additional material examined. - india: Paratypes 2 9 ,
Assam 5000ft. Shillong, 2.vi.l924, Fletcher leg.; Khasis,
5000 ft. Shillong, ló\ 25.V.1923, Fletcher coll.; Assam,
Shillong, 1 9, H.M. Parish (bmnh).
Heterothera taigana taigana (Djakonov)
Cidaria taigana Djakonov, 1926: 27. Holotype 3, Russia:
Sajan Mountains. Kasyr, E. Minusinsk, 7.viii.[19]24, L. &
I. Koshantschikov (Djakonov 1926: 27) [not examined].
Asaphodes taigana; Viidalepp 1980: 65.
Viidaleppia taigana; Inoue 1982: 283.
Heterothera taigana; Choi 1 997: 311.
Diagnosis
H. taigana is characterized by the dark ochreous or
blackish ground colour of wings, small blackish dot at
the dorsum and the costally smooth waving post-
medial line of forewing. In the male genitalia the un-
cus, anellus lobe, costa, and sacculus are characteris-
tic. The females of taigana are distinguished by the
very large horn-like lamella postvaginalis.
Redescription
Antenna of male bipectinate, with short pectina-
tions. Frons blackish. Labial palp about 1.5 times as
long as eye diameter. Legs blackish with white tibial
joints. Wingspan 27-33 mm. Forewing ochreous
brown; basal line blackish, forming two vertical lines,
waved, slanted; antemedial line waved, medially in-
dented; postmedial line black, medially round, bul-
ged; central fascia even in width, with black discoidal
dot; subterminal line blackish, round; dorsum be-
tween basal and antemedial lines distinct with black
dot. Hindwing blackish; discoidal dot black, small;
postmedial line blackish.
Male genitalia. - Uncus sclerotized, twice as long as
tegumen, basally tapered, medially bent. Tegumen
dome-shaped. Saccus shallow, broad. Anellus lobe
large, horizontally expanded, rod-shaped, with long
hairs on dorsal part. Valva sclerotized in general, dis-
toventrally membranous; costa largely sclerotized,
with distal end blunted; sacculus large, triangular,
ventrally very slender, distoventrally with sharply
pointed process. Aedeagus long, slender, sclerotized,
distally scobinated; vesica tubular with small divertic-
ula; cornuti mixed with large and small spines.
Female genitalia. - 8th segment sclerotized. Anteri-
or apophysis absent. Posterior apophysis long, slender.
Lamella postvaginalis with large horn-like sclerotized
process; lamella antevaginalis plate-like, sclerotized.
Ductus bursae short, broad, wall with sclerotized
processes. Corpus bursae slightly slender or ovate.
Distribution
Russia (eastern Siberia).
Remarks
Djakonov (1926) noted its closeness to the ßrma-
ta-group in the wing pattern.
Material examined. - Russia: Irkutskaja obi., Hamar-Da-
ban, taiga, Meteorolog st. 1450 m, 63 1 9, 25-27.vii.1984,
ad. lue. Mikkola & Viitasaari; Magadan obi., Upper Kolyma
r. 62°N 149°40'E, 600 m scree slope ad. lue, lo*, I6.vii.
1987, K. Mikkola; 59°10'N 150°E, Magad. obi., Ohkotsk
coast, 100 Km W. Magadan, 1 cT 29, 4-6.viii.1989, K.
Mikkola (zmh); Baikal, SW coast, Shumilikha river, 13,
10.viii.1972, (ZMI).
Heterothera taigana sounkeana (Matsumura)
Cidaria {Thera) sounkeana Matsumura, 1927: 184. Holo-
type 3 , japan: Sounkei, Hokkaido (Mt. Daisetsu), 9.viii.
1926, Matsumura leg. [not examined].
Thera kurilaria Bryk, 1942: 70. [Synonymized by Inoue
1955:72].
Thera kurilaria ecce Bryk, 1942:71. [Synonymized by Inoue
1955:72].
Cidaria hospes Djakonov, 1955: 318.
Asaphodes sounkeana; Viidalepp 1980: 65.
Viidaleppia taigana sounkeana; Inoue 1982: 283.
Thera taigana ishizukai Inoue, 1955: 72. Holotype 3,
japan: Mt. Jonen (2500m), Japan Alps, Nagano Pref.,
26.vii.1951, H. Inoue (bmnh) [examined], syn. n.
Diagnosis
This subspecies is separated from the nominal sub-
species by the presence of the signum in the corpus
bursae.
Distribution
The Russian Far East, Japan.
Remarks
Inoue (1955) noted that Thera kurilaria Bryk
might be the same species as sounkeana Matsumura on
39
Tijdschrift voor Entomologie, volume mi, 1998
Figs. 52-55.
Female genitalia of Heterothe-
ra. — 52, H. consimilis; 53, H.
êtes; 54, H. serrarla; 55, H. ku-
renzovi.
the basis of the length of fotewing and the structure of
male genitalia. He (1977, 1982) listed two subspecies
of taigana in Japan, sounkeana and ishizukai.
The subspecies sounkeana has the derived feature in
the female genitalia; the presence of one signum
patch in corpus bursae. This subspecies ranges from
Amur through Kurile Islands to Japan.
In the original description of ishizukai, Inoue
(1955: 72) wrote the differences between these two
subspecies: "larger, and a little lighter in colour and
postmedian line of forewing more weakly incurved in
cellule 6, discal fleck a little less heavier than soun-
keana." I have checked 3 type specimens of ishizukai,
and the subspecific rank of ishizukai compared to
sounkeana was very doubtful. Thus, I synonymized
ishizukai with sounkeana.
Biology
Pinus pumila Regel is known as a host plant in
Japan (Sato & Nakajima 1987).
Material examined. - Russia: Kemorovo obi. Gornaya,
Shoriya, 1$, 24.viii.1965, Kononenko; Kurilid, Itunup,
1 a, 25.vii.1979, V. Junno; Sahhalin, Birjusinka, L., 1 9 , 7-
10.vii.1975, Viidalepp ; Kuriilid, Kunasir, Tretjakovo, 1 $,
22- 24.vii.1980, Pototski, Ruben & Veldre (zbi). - japan:
Mt. Daisetsu, Hokkaido, Id 2?, 29.vii.1978, Y. Kishida;
Kamikawa Mt. Daisetsu, Hokkaido, Id, 20- 21.viii.1993,
H. Kobayashi (ky); Mt. Mitsumata Range 2560 m, 1 ex.,
4.viii.l954, T. Haruta (zmh).
40
CHOI: Systematics of Heterothera
Heterothera serrarla (Lienig)
(fig. 54)
Cidaria serrarla Lienig, 1846: 200. Holotype baltic
(latvia). [not examined]
Gnophos sen-aria Herrich-Schäffer, 1847: 72.
Cidaria llenlglarla Lederer, 1853: 183 [Synonymized by
Staudinger & Wocke 1871: 184].
Cidaria serrarla Zeller; Staudinger & Wocke 1871: 184,
Prout 1914: 219 [Incorrect authorship].
Melanippe ziczaecata Schöyen, 1875: 145. [Synonymized by
Prout 1914:219].
Larentia serrarla ab. continua Strand, 1903: 16.
Larentla serrarla ab. albida Stichel, 1911: 86.
Larentia serrarla ab. spania Stichel, 1 9 1 1 : 86.
Asaphodes serrarla, Viidalepp 1977: 576.
Vlldaleppla serrarla; Inoue 1986: 61.
Heterothera serrarla; Choi 1997: 311.
Diagnosis
H. serrarìa is characterized by the whitish ground
colour, blackish dots between basal and antemedial
lines, medially broken central fascia, white and scal-
loped subterminal line of forewing and the blackish
and scalloped subterminal line of hindwing. In the fe-
male genitalia, the antrum and very short and heavily
sclerotized ductus bursae are distinguishing features.
This species is very similar to the following two species,
H, serrataria and kurenzovi, but differs in the termen
of hindwing and the sterigma of female genitalia.
Redescription
Antenna of male bipectinate, with long pectina-
tions. Frons whitish, round. Labial palp short, less
than eye diameter. Metathorax dorsally white with
blackish tufts. Wingspan 21-26 mm in male, 27-29
mm in female. Forewing ground colour whitish; basal
line black, dentate; area between basal and antemedial
line with blackish vertical band; antemedial line medi-
ally slightly indented; postmedial line costally waved,
medially bulged; central fascia blackish, sometimes
medially missing or reduced, discoidal dot large,
blackish; termen blackish, subterminal line white,
waving. Hindwing whitish; discoidal dot blackish; ter-
men black with whitish, waving subterminal line. 1st
tergal sclerite minute.
Male genitalia. - Uncus thin, shorter than tegu-
men, medially slightly bent. Tegumen bell-shaped.
Saccus medially indented, broad. Juxta broad, sclero-
tized. Anellus lobe small round, sclerotized, very short
hairs on top. Valva distally tapered; costa sclerotized,
with medial expansion; sacculus well developed, dis-
toventrally a large process. Aedeagus cylindrical, scle-
rotized; vesica tubular, cornuti large spinular.
Female genitalia. - Anterior apophysis absent, pos-
terior apophysis long, slender. Lamella postvaginalis
semi-circular covered with minute spines, lamella an-
tevaginalis sclerotized, V-shaped with medially in-
vaginated. Ductus bursae very short. Corpus bursae
large, posteriorly strongly sclerotized, with sclerotized
striations at anterior part. Signum absent.
Distribution
Northern Europe to eastern Siberia.
Remarks
The H. serrar ia-group, comprising three species,
H. serrarla, serrataria and kurenzovi, is characterized
by the wing pattern and the female genitalia. Among
these species, the latter two species are sympatric in
the Russian Far East. The species fly concurrently
during July in southern Ussuri and Habarovsk. The
species kurenzovi can be distinguished by more black-
ish coloration of the wing pattern elements and espe-
cially, by the genitalia of both sexes. The distal part of
the aedeagus in serrataria is sclerotized and the cornu-
ti are more or less small spinular processes, but the
aedeagus of kurenzovi is distally sclerotized and also
several spines are present at the vesica and the cornu-
ti are the mixture of large and small spines in the vesi-
ca. In the female genitalia, the lamella antevaginalis of
serrataria is a pair of triangular processes, whereas that
of kurenzovi is a pair of rectangular processes.
Biology
Probably Picea is the host plant, and the species hi-
bernates in the larval stage (Mikkola et al. 1985).
Material examined. - Finland: Seinäjoki.l â 1 9, 30.vi.
1931;Porvoo,Aminsby, let 1 9 , 23.vi.1959, 24.vi.1970, E.
Suomalainen. - Russia: Murmansk district, 65 Km S. Mon-
chegorsk, 3cT 2$, 19.vii.1993, A. Lvovsky; Murmansk dis-
trict, Luvenga, White sea shore, 23 1 9 , 29.vi.1989, M. Ko-
zlov; Petropolis, 1 â 1 9 , Coll. Duske; S-Ural, Cheliabinsk
dist., Iremel mountain reserve, 900-1300 m, 7(5", 23-
27.vi.1996, K. Nupponen, J.-P. Kaitila, J. Junnilainen & M.
Ahola; Altai Sibr., 1 9 , Coll. Winter; Irkutskaja obi.,
Hamar-Daban, taiga, Meteorolog st. 1450 m, 13Ó* 49, 25-
27.vii.1984, ad. lue. Mikkola & Viitasaari; Buryatia,
53°13'N 109°19'E, Barguzin valley 1000m, Ust- Barguzin-
Yambulrd. taiga, 5d , 13.vii.1996, Jalava& Kullberg (zmh).
Heterothera serrataria (Prout) comb. n.
(figs. 25, 36)
Cidaria (Thera) serrarla serrataria F 'rout, 1914: 219. Holo-
type 6 , Russian far east: Kasakewitsch, Ussuri, in Coll.
Püngeler (Prout 1914: 219) [not examined].
Cidaria ( Thera) serrataria; Prout 1938: 115.
Asaphodes serrataria; Viidalepp 1977: 576.
Vlldaleppla serrataria; Inoue 1986: 60.
Diagnosis
H. serrataria is very similar to H. serrarla, but dif-
fers in the complete central fascia, the termen of hind-
wing without blackish subterminal line, the digiti-
form anellus lobe, the deep saccus and the lamella
post- and antevaginalis.
41
Tijdschrift voor Entomologie, volume 141, 1998
Redescription
Antenna of male bipectinate, with long pectina-
tions. Frons brownish black. Labial palp short, less
than eye diameter. Wingspan 24-28 mm. Forewing
ground colour whitish; basal line blackish, dentate;
blackish vertical dots between basal and antemedial
lines; antemedial line costally largely toothed, medial-
ly indented; postmedial line costally toothed with one
deep invagination, medially bulged; central fascia
blackish, with blackish discoidal dot; termen blackish;
subterminal line waving, whitish. Hindwing whitish;
basal blackish, with weak discoidal dot; postmedial
line blackish. 1st tergal sclerite moderate projection.
Male genitalia. — Uncus short, sclerotized, basally
tapered. Tegumen bell-shaped. Saccus broad, medial-
ly indented. Juxta broad, basally round. Anellus lobe
digitiform, small hairs on top. Valva slim, distoven-
trally expanded; costa sclerotized, medially and distal-
ly expanded; sacculus triangular, sclerotized, dis-
toventrally with process. Aedeagus cylindrical, distally
strongly sclerotized; vesica tubular, cornuti spinular.
Female genitalia. - Anterior apophysis absent, pos-
terior apophysis long, slender. Lamella postvaginalis
large round processes with toothed surfaces, lamella
antevaginalis broad, V-shaped, medially hollowed,
covered with spines. Ductus bursae short, heavily
sclerotized. Corpus bursae large, strongly sclerotized,
with broad opening.
Distribution
The Russian Far East, Japan.
Remarks
Inoue (1986) listed this species under Viidaleppia
on the basis of the genitalia. However, he was uncer-
tain of its generic status because the species showed
slender pectinations in male antenna which was simi-
lar to Pennithera, and single aréole of forewing which
was exceptional in Thera- group.
Jaros et al. (1992) recorded the species H. serrarla
from the Korean Peninsula. However, the record is
very suspicious based on the distribution and I suppose
that the specimen might be H. serrataria or kurenzovi.
Material examined. - Russia: S-Ussuri, Id" 19,9-
12.vii.1976, Tsugujevski rj. dvnts, Berjzovoi stats, drl.
Metsavir, Viidalepp, Ruben & Vasjurin; Habarovsk, Nel-
ma, 29, 16-26.vii.1977, Viidalepp, Laanetu & Talve (zbi);
Pompejefka, Amur, 2c? 1 9 (bmnh). - japan: Mt. Peipan,
Asahikawa C. 1 ó\ 12.vii.83, Y. Kusunoki (bmnh).
Heterothera kurenzovi Choi,Viidalepp & Vasjurin sp.n.
(figs. 14, 26, 37, 55)
Pennithera kurenzovi Viidalepp 1996: 23. [nomen nudum]
Type material. - Holotype 3 , Russian far east:
Habarovsk, Nelma, l6-26.vii.1977, Viidalepp, Laa-
netu & Talve leg. (zbi). Para type 1 5 S-Ussuri, Tsug-
ujevski rj. dvnts, Berjzovoi stats, drl, Metsavir, 9-
12.vii.1976, Viidalepp, Ruben & Vasjurin (zbi).
Diagnosis
This species is very similar to H. serrataria, but dif-
fers in the more or less dark wing elements and medi-
ally broken central fascia. The long spinular processes
at the distal part of the aedeagus, the very large spin-
ular cornuti and the sterigma of the female genitalia
are characteristic.
Description
Antenna of male bipectinate, with long pectina-
tions. Frons brownish black. Labial palp short, less
than eye diameter. Metathorax dorsally white with
blackish tufts. Wingspan 26-28 mm. Forewing
ground colour whitish; basal part blackish, basal line
dentate; next to basal line are blackish, vertical dots;
antemedial line costally and dorsally projected in-
ward, medially indented; postmedial line generally
waved, medially bulged; central fascia blackish, with
strong blackish discoidal dot, medially broken; ter-
men with round blackish, vertical band, subterminal
line white, waved. Hindwing whitish, discoidal dot
very weak; termen weak blackish spots. Metathorax
whitish. 1st tergal sclerite with moderate process.
Male genitalia. - Uncus short, sclerotized, basally
tapered. Tegumen bell-shaped. Saccus broad, medial-
ly indented. Juxta broad, sclerotized. Anellus lobe
digitiform, apically with very short hairs. Valva basal-
ly broad, distally reduced; costa slender, sclerotized,
medially largely expanded; sacculus triangular, sclero-
tized, distoventrally with a claw-like process. Aedea-
gus cylindrical, distally with spinular processes, cor-
nuti large, spinular, scattered.
Female genitalia. - Anterior apophysis absent, pos-
terior apophyses long, slender. Lamella postvaginalis
semi- circular processes with dentate surfaces, lamella
antevaginalis large plate-like structure densely covered
with sclerotized processes. Ductus bursae very short,
sclerotized. Corpus bursae large, opening with strong-
ly sclerotized, anterior part with sclerotized striations.
Distribution
The Russian Far East (Primorye, Sikhote Alin
Mts.), Sakhalin, N. Japan (Viidalepp 1996).
Remarks
Originally, the new species was described a decade
ago by Viidalepp and Vasjurin on the basis of the ma-
terial which is now preserved in the Institute of Biol-
ogy and Pedology, Vladivostok. However, unfortu-
nately this description had not yet been published.
During my preparation of Heterothera, Dr. Viidalepp
42
CHOI: Systematic! of Heterothera
kindly provided the material. Thus, here, we are de-
scribing this species jointly.
Heterothera firmata (Hübner)
(fig. 24)
Geometra firmata Hübner, 1822: 515. Holotype Europe
[not examined].
Thera consobrinata Curtis, 1834: 519 [Synonymized by
Prout 1914:219].
Thera simulata Stephens, 1831: 271 nee. Hübner, 1822
[Synonymized by Staudinger & Wocke 1871: 184].
Chesias ulicata Rambur 1834: 394; Staudinger & Wocke
1871: 184, Prout 1914:219.
Larentia firmata; Herrich-Schaffer 1847: 171.
Cidaria firmata; Staudinger & Wocke 1871: 184.
Thera firmata; Viidalepp 1977: 576.
Thera firmata ab. approximata Lempke, 1950: 163.
Thera firmata ab. brunneofasciata Lempke, 1950: 163.
Thera firmata ab. grisescens Lempke, 1950: 163.
Thera firmata ab. interrupta Lempke, 1950: 163.
Thera firmata ab. purpureobrunnea Cockayne, 1952: 267.
Asaphodes {Pennithera) firmata; Viidalepp 1980: 70.
Heterothera firmata; Choi 1997: 311.
Diagnosis
The wing pattern elements and the wing colour of
H. firmata are somewhat similar to Thera obeliscata.
However, firmata can be easily distinguished by the
bipectinate male antenna, strongly indented anteme-
dial line and the blackish discoidaJ line that is fused to
the costal part of the antemedial line of forewing.
Redescription
Antenna of male bipectinate, with long pectina-
tions. Frons yellowish brown. Labial palp long, double
eye diameter. Legs yellowish brown, with pale tibial
joints. Wingspan 24-30 mm. Forewing yellowish;
basal line waved, occasionally costally strongly bulged;
antemedial line costally blackish, medially deeply in-
dented; postmedial line costally blackish, medially
bulged; central fascia dark yellowish, dorsally reduced,
blackish discoidal dot; subterminal line whitish, waved.
Hindwing yellowish white, postmedial line very weak.
Male genitalia. - Uncus strongly sclerotized, medi-
ally bent, short being less than tegumen height. Tegu-
men triangular, shorter than vinculum. Saccus shal-
low, round. Juxta well developed, sclerotized. Anellus
lobes digitiform, apically expanded and hairs present.
Valva midventrally expanded; costa strongly sclero-
tized, with median and distal expansions; sacculus
sclerotized, with distoventral process. Aedeagus cylin-
drical, sclerotized; vesica tubular, basally one patch of
spinular cornuti present.
Female genitalia. - Anterior apophysis about 1/5
length of posterior apophysis. Antrum simple. Duc-
tus bursae long, strongly sclerotized, basally thick,
distally reduced. Corpus bursae ovate, small.
Distribution
Europe.
Remarks
Pierce (1914) first pointed out that the genitalia of
H. firmata are different from Thera s.s. The species
was transferred from Thera to Pennithera by Viidalepp
(1980), and subsequently it was again moved from
Pennithera to Heterothera (Choi 1997). The male and
female genitalia of firmata., however, are somewhat
different from the typical shape of Heterothera and
the cladistic reanalysis of Thera and related genera
(Choi unpublished) proposes that the species firmata
is separated from Heterothera. Therefore, the generic
position of the species is unclear.
Biology
The species is sometimes bivoltine in England and
southern Europe (e.g. Bulgaria), although it is univol-
tine in northern Europe. Pinus is known as a host
plant (Mikkola et al. 1985).
Material examined. - Finland: Porvoo, 1 S , 31.viii.1952,
Seppänen; Sibbo, 1 â , 1 l.viii. 1983, A. Albrecht; Tvärminne
Biol. St., 19, 13.ix.1951, E. Suomalainen; Helsinki.l 9,
25.viii.1971, A. Albrecht(zMH). - Austria: Meran, Tirol,
1 9, coll. Winter (zmh). - Hungary: Sopron, Tacsi drok,
lo\ 31.viii.1978, Ronkay; Sopron, Kovas d., 10* 19,
16. ix. 1949, Ambrus; Peszerpuszta, Hungaria cent., 19,
24.ix.1949, Dr. Issekutz (hnhm). - Bulgaria: Rhodope Mt.
Lukovitza motel near Assenovgrad, 450 m, lo*, 31.vii.1985,
20*, 3.x. 1986, S. Beschkov; S.W. Bulg. Kresnensko defile,
Station Stara Kresna, 200 m, lo", 17.X.1987, S. Beschkov;
Troyanska Stara planina mount, hut Dermenka, 1533 m,
1 9, 21. ix. 1987, S. Beschkov (bnhm).
Thera cyphoschema Prout incertae sedis
(figs. 40,41,56)
Cidaria {Thera) cyphoschema Prout, 1926: 312. Holotype o*,
N.E. burma: Htawgaw, 6000 ft. March 1923, A. E. Swann
leg. (bmnh) [examined].
Thera cyphoschema; Prout, 1941: 323.
Cidaria atrinotata Joannis, 1929: 485. Holotype o", Viet-
nam: Hoang su phi, Tonkin in Paris Museum (colour
transparency in bmnh, [examined]), syn. n.
Cidaria atrinotata reducta Joannis, 1929: 486. syn. n.
Diagnosis
This species is similar to H. mussooriensis in the wing
pattern, but differs in the short pectinations of the
male antenna and the relatively short and thick uncus,
round saccus, unsclerotized costa of valva, the presence
of diverticulum of vesica and the absence of cornutus.
Redescription
Antenna of male bipectinate, with short pectination.
Frons reddish and white scales. Labial palp brownish
and white, long about twice the eye diameter. Legs
43
Tijdschrift voor Entomologie, volume ui, 1998
Figs. 56-57.
Female genitalia of Thera s.l.
T. cyphoschema; 57, T. exa:
Scale bar= 1 mm.
56,
brownish and white scales, with white tibial joints.
Wingspan 26 mm. Forewing ground colour greyish;
basal line blackish, slanted; antemedial line costally
blackish, medially indented; postmedial line black,
medially bulged; central fascia thin, medially reduced,
blackish on veins; dorsum with short blackish line at
base. Hindwing whitish, discoidal dot blackish, post-
medial line very weak.
Male genitalia. - Uncus thick, relatively short less
than half of tegumen, sclerotized. Tegumen dome-
shaped. Juxta broad, sclerotized; anellus lobe digiti-
nomi, medially and apically with long hairs. Saccus
round, medially concave. Valva generally membra-
nous; costa membranous, distally slightly projected
upward; sacculus triangular, basally sclerotized, with
a distal large spinular projection. Aedeagus slender,
sclerotized; vesica tubular with one large lateral diver-
ticulum; cornutus absent.
Female genitalia. — Anterior apophysis half of pos-
terior apophysis. Sterigma simple, laterally with thin,
sclerotized stripes. Antrum large, funnel-shaped.
Ductus bursae with colliculum in middle. Corpus
bursae large, subspherical, signum absent.
Distribution
Burma, S. China, N. Vietnam.
Remarks
The species is similar to Heterothera in the bipecti-
nate male antenna, the wing pattern and the male
genitalia, but it is very different from Heterothera in
the female genitalia, particularly the funnel-shaped
antrum and the presence of a colliculum. Thus, the
inclusion of the species in Heterothera is equivocal.
Additional material examined. — burma: Paratypes 3 c? ,
Htawgaw, N.E. Burma, 6000 ft. IV- V.1923, A.E. Swann.;
Htawgaw, N.E. Burma, 6000 ft, Id 3?, IV-V.1923, A.E.
Swann. — Vietnam: Paratype 1 â , Hoang su phi, Tonkin. -
china: Tseku (Tse-Kou), Prov. Yunnan, 3d", 1895, 2â ,
1898, 5ó\ 1900, R.P.J. Dubemard (bmnh).
Thera exangulata (Warren) incertae sedis
(figs. 42, 43, 57)
Perizoma? exangulata Warren 1909: 127. Holotype ?, n.
india: Srinagar, Kashmir 7000ft, collection]. Ward.
20.VI.[19]04 (bmnh) [examined].
Cidaria {Thera) exangulata P 'rout 1938: 113.
Thera exangulata; Prout 1941: 323.
44
CHOI: Systematics of Heterotbera
Diagnosis
The species can be distinguished by the black and
medially reduced central fascia and white scalloped
subterminal line of forewing. This species is similar to
Cidaria deletaria Hampson in the male genitalia (the
slender and sclerotized costa), but differs in the wing
pattern elements.
Redescription
Antenna of male filiform. Frons smooth, mixed
with ochreous and brownish scales. Labial palp short.
Legs dark brownish, with ochreous tarsal joints.
Wingspan 32 mm. Forewing basal part dark brown-
ish, slanted; antemedial line costally expanded, slant-
ed; postmedial line medially dentate, bulged; central
fascia blackish, costally pale brownish, medially re-
duced, with long black discoidal dot. Termen brown-
ish, medially paler; subterminal line whitish, scallop-
ing; apical streak brownish. Hindwing whitish; basal
part tinged with blackish, with blackish discoidal dot;
postmedial line blackish; termen dorsally slightly
tinged with black.
Male genitalia. - Uncus about twice as long as
tegumen, basally tapering. Tegumen dome-shaped.
Saccus round, medially expanded. Juxta broad; anel-
lus lobe long, rod-shaped, dorsally with long hairs.
Transtilla sclerotized, thin, round. Valva slender; cos-
ta long, slender, distally with long sclerotized hairs;
sacculus indistinct. Aedeagus cylindrical, sclerotized,
ductus ejaculatorius distally sclerotized; vesica large
sac-like, cornutus absent.
Female genitalia. - Papillae anales simple, sclero-
tized. 8th segment slightly sclerotized. Anterior apoph-
ysis about 3/5 length of posterior apophysis. Antrum
broad, funnel-shaped. Ductus bursae short, with col-
liculum. Corpus bursae very large, subovate, posteriorly
with many striations; signum a long thread-like process.
Distribution
N. India.
Pennithera distractata (Sterneck) comb. n.
(figs. 44, 45)
Thera distractata Sterneck, 1928: 152. Holotype 9, china:
Wassukou (Wassekou), Prov. Sichuan [not examined].
Diagnosis
The species is similar to H. serraria and H. ser-
rataria in the wing pattern elements, but it can be dis-
tinguished by the short pectinations of male antenna,
its wider central fascia and the male genitalia. The
well developed bilobed anellus lobe and the medial
process at the costa of valva in the male genitalia are
characteristic.
Description
Antenna of male bipectinate, with short pectina-
tions. Frons with white and blackish scales. Labial
palp moderate about 1.5 times the eye diameter. Legs
with whitish tibial joints. Wingspan 25 mm in male.
Forewing ground colour whitish; basal line blackish,
waved; area between basal and antemedial lines tinged
with blackish, bar-shaped band; antemedial line
waved; postmedial line costally dentate, invaginated,
medially slightly bulged; central fascia blackish, rela-
tively constant in width, with black discoidal dot; ter-
men blackish. Hindwing whitish; discoidal dot small
blackish, with blackish postmedial line; termen with
black, waving subterminal line; fringe black.
Male genitalia. - Uncus long, slender. Tegumen
small, dome-shaped. Juxta basally invaginated; anellus
lobe bilobed with club-shaped large lobe and densely
hairy small lobe, apical part of large lobes with long
hairs. Saccus sclerotized, round with medial projec-
tion. Valva slender; costa sclerotized, with a medial
triangular process. Aedeagus slender, medially bent,
distally slightly scobinated; cornutus small process.
Female genitalia. - Unknown.
Distribution
S.W. China.
Remarks
Warren (1909) described the species under Peri-
zoma, but Prout (1938 & 1941) listed it under Thera.
The wing pattern and especially the female genitalia
of the species (funnel-shaped antrum, the presence of
colliculum and the thread-like signum) are different
from Thera s.l. The male genitalia of exangulata are
similar to Cidaria deletaria Hampson, whereas the
antrum of female genitalia is similar to T. cyphosche-
ma. The systematic position of the species remains un-
clear.
Remarks
The species has never been reported since its origi-
nal description (Sterneck 1928). The identification of
the species is based on the original description and
drawing.
The combination with Pennithera is based on the
following characters: the relatively smaller tegumen,
the triangular process at the middle of the costa and
the indistinct sacculus of the male genitalia. The
bilobed anellus lobes of the male genitalia are very
characteristic.
Additional material examined. - india: Kashmir, 7000 ft,
2d\ 24.vi.; Mussu Hills, 2$ (bmnh).
Material examined. - china: 1 ô A-tun-tse, (Nord- Yun-
nan), Obere Hohe, ca. 4500 m, 13.vii. 1937, H. Hone (zfmk).
45
Tijdschrift voor Entomologie, volume i4i, 1998
Acknowledgements
I thank Dr. F. Rindge (amnh), Dr. M. Scoble, Miss
K. Buckmaster (bmnh), Dr. A. Popov (bnhm), Dr. L.
Ronkay (hnhm), Mr. K. Yazaki (Tokyo), Mr. M.
Fibiger (Copenhagen), Mr. B. Gustafsson (snhm),
Dr. J. Viidalepp (zbi), Dr. D. Stüning (zfmk), and
Dr. A. Hausmann (zsm) for providing the material for
this study. Thanks are due to Mr. R. Tyynelä and A.
Rusanen for taking the photographs. Dr. Malcolm
Scoble (London) and Dr. Kauri Mikkola (Helsinki)
made helpful suggestions on an earlier draft. Dr. Jaan
Viidalepp (Tartu) helped in many ways to this manu-
script. The kind assistance in the editorial process and
final publishing of the paper given by Dr. Erik J. van
Nieukerken (Leiden) is also appreciated. The study
was financially supported by the cimo, Finland.
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Sato, R. & H. Nakajima, 1987. Geometridae. - In S. Sugi
(ed.), Larvae of larger moths in Japan, pp. 31-1 12. — Ko-
dansha, Tokyo.
46
CHOI: Systematics ofHeterothera
Schöyen, W. M., 1875. Fortegnelse over Sommerfugle. —
Nytt Magasin for Naturvidenskabene 21: 139-146.
Staudinger, O. & M. Wocke, 1871. Catalog der Lepi-
dopteren des Europaeischen Faunengebiets. — Dresden.
426 pp.
Stephens, J. F., 1829-1831. Illustrations of British Ento-
mology. Haustellata, 3. - Baldwin and Cradock, London.
333 pp.
Sterneck, J., 1928. Die Schmetterlinge der Stötznerschen
Ausbeute. Geometridae, Spanner. - Deutsche Entomolo-
gische Zeitschrift, Iris 42: 131-183.
Stichel, H., 1911. Zweiter Beitrag zur nordischen Schmet-
terlinges fauna und anknüpfende Bemerkungen. - Ber-
liner Entomologische Zeitschrift 56: 33-104.
Strand, E., 1903. Neue norwegische Schmetterlingsformen.
Archiv for Mathematik of Naturvidenskab 25(3: 9): 1-24.
Viidalepp, J., 1977. A list of the Geometridae (Lepidoptera)
of the USSR. Part II. - Entomologiceskoe Obozrenie
16(3): 564-576. [In Russian]
Viidalepp, J., 1980. Geometrid moths of the genus Thera
Stph. in the fauna of the USSR (Lepidoptera). - Tartu
Riikliku Ulikooli Toimitised 13(516): 54-84. [In Russian
with English summary]
Viidalepp, J., 1996. Checklist of the Geometridae (Lepi-
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Warren, W., 1888. On Lepidoptera collected by Major Yer-
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Warren, W., 1909. New species of Thyrididae, Uraniidae,
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Received: 20 October 1997
Accepted: 13 January 1998
47
Tijdschrift voor Entomologie, volume hi, 1998
BOOK REVIEWS: ORTHOPTERA SOUNDS
D.R. Ragge & W.J. Reynolds, 1998. The songs of the
Grasshoppers and Crickets of Western Europe. - Harley
Books, Colchester. 591 pp., 3 colour plates, 24,4 x 17,2
cm, clothbound. [isbn 0-946589-49-6]. Price £ 65.00
excl. p&p.
D.R. Ragge & W.J. Reynolds, 1998. A sound guide to the
Grasshoppers and Crickets of Western Europe. - Harley
Books, Colchester. Jewel box with 2 cd's, 120 minutes,
booklet 20 pp.. [isbn 0-946589-50-X]. Price £ 25.00
excl. p&p.
The editors received the book and the CD-booklet
for review, but for completeness sake I'll also include
the cd's themselves here. Below I also take the oppor-
tunity to briefly review the other recent European
books and cd's containing Orthoptera sounds. Being
an editor and co-author of the Dutch book and CD
myself, I'll try to be as objective as possible.
The book and cd's by Ragge & Reynolds are the
first to treat the songs of the majority of Western Eu-
ropean singing Orthoptera in full detail (Heller made
a more complete review of busch-crickets only) . The
book describes the songs of 170 species occurring in
western Europe and Scandinavia, including Finland,
Austria, Italy and the western Mediterranean islands.
Each description comprises the full Latin name and
original citation; vernacular names of the major lan-
guages of the countries covered; a complete set of ref-
erences to earlier song descriptions and recordings; a
paragraph on recognition; full description of the song
in its various types (calling song, courtship, rivalry
song etc.) and a paragraph on distribution. The songs
are illustrated by ca. 1600 oscillograms. All the songs
are also included in the accompanying cd's, in rela-
tively short fragments (ca 25 seconds each). Three
colour plates give an impression of the diversity of the
singing Orthoptera in western Europe.
The introduction gives many technical details on
recording and other acoustic methods,, and provides
a general introduction and reviews of sound produc-
tion in Orthoptera and nature and function of songs.
In the introduction the used nomenclature is ex-
plained for difficult cases. A final introductory chap-
ter discusses the value of songs in taxonomy and iden-
tification.
The book is a major contribution to Orthoptera
systematics, it not only describes the songs of many
species, but also includes the many taxonomie results
which were published by these and other authors in
earlier papers. The careful study of the songs has in
many cases lead to a better understanding of complex
groups, such as the Chorthippus biguttulus group. The
authors could reduce many dubious very local forms
into synonyms of more widespread species. The cd's,
oscillograms and key are very handy tools for the
identification on the basis of sound. I like the inclu-
sion of the similar sounds of the four larger cicadas,
two toads and five bird-species. I only wonder why
the toad Bufo calamita is not included: here in Hol-
land it's song is often confused with Grillotalpa song.
The species included are apparently chosen on the
pragmatic basis of material available. They represent
all common and most rarer species, but nowhere the
authors explain what proportion of Orthopteran
species they actually represent. Also missing is a list of
the not-singing groups which are not covered. In
some groups the selection seems rather artificial, such
as the Locustinae, where Psophus stridulus and Bryode-
ma tuberculatum are treated, but the similar Oedipo-
da-species, which also make sounds, not. This is the
only point of criticism on an otherwise very useful
and important book and CD-set, that I'll recommend
warmly for all those interested in Orthoptera.
Other titles
H. Bellmann, 1993a. Heuschrecken beobachten, bestim-
men. - Naturbuch Verlag, Augsburg [Second revised edi-
tion] ,349 pp. [isbn 3-89440-028-5]. [in German]
H. Bellmann, 1993b. Die Stimmen der heimischen Heu-
schrecken. - Weltbild Verlag Augsburg, audio-cd.
H. Bellmann & G. Luquet, 1995. Guide des sauterelles,
grillons et criquets d'Europe occidentale. - Delachaux et
Niestlé, Lausanne, 383 pp. [isbn 2-603-00974-5]. Price
appr. NLG 80. [In French]
F.-R. Bonnet, 1995. Guide sonore des sauterelles, grillons et
criquets d'Europe occidentale. - Delachaux et Niestlé,
Lausanne, Audio-CD. [isbn 2-603-00975-3]. Price appr.
NLG 30.
R.M.J.C. Kleukers, E.J. van Nieukerken, B. Odé, L.P.M.
Willemse & W.K.R.E. van Wingerden, 1997. De sprink-
hanen en krekels van Nederland (Orthoptera). - Neder-
landse Fauna 1. Nationaal Natuurhistorisch Museum,
knnv Uitgeverij & Eis-Nederland, Leiden. 408 pp., 14
plates, [isbn 90-501 1-100-9]. price nlg 82.50 including
cd, excl. p&p. [In Dutch, extensive English summaries]
[end 1998: almost sold]
B. Odé, 1997. De zingende sprinkhanen en krekels van de
Benelux. Audio-CD, included with book Kleukers et al..
Both Bellmann guides are new editions of his earli-
er fieldguide in German, of which also an English
translation exist. They treat many species which were
not earlier included, and therefore extent the region of
use enormously. With the magnificent photographs
48
continued on page 64
Hans R. FEIJEN
National Museum of Natural History, Leiden
TELEOPSIS RONDANI (DIPTERA, DIOPSIDAE):
GENERIC REVIEW AND THE FERRUGINEA GROUP
FROM SRI LANKA
Feijen, H. R., 1998. Teleopsis Rondani (Diptera, Diopsidae): generic review and the ferruginea
group from Sri Lanka. - Tijdschrift voor Entomologie 141: 49-63, figs. 1-33, table 1. [issn
0040-7496]. Published 30 November 1998.
Teleopsis Rondani is defined. A catalogue is presented for the 1 9 species now recognized, while
the systematic history of the genus is discussed. New synonymies are proposed and several old
synonymies rejected. Various characters of taxonomie importance are discussed. For all species,
data are provided on two quantitative characters: ratio eye span/body length and ratio
length/width of front femur. Both characters show a correlated interspecific variation from
strong sexual dimorphy to homomorphy. A key is provided to the five Diopsidae known to oc-
cur in Sri Lanka. From Sri Lanka Teleopsis ferruginea Roder is redescribed, while two new
species are described: Teleopsis krombeini sp. n. and Teleopsis maculata sp. n.
H. R Feijen, P. O. Box 639, Thimphu, Bhutan or (permanent) P. O. Box 2068, 2301 CB Leiden,
The Netherlands
Keywords. — Diptera; Diopsidae; Teleopsis; revision; new species; redescription; catalogue; sexual
dimorphy; Oriental; Sri Lanka.
The genus Teleopsis was created by Rondani in
1875. As type-species, Rondani designated Diopsis
sykesii Gray, though Westwood is now commonly
considered the author of this species. No formal diag-
nosis was given. Only in his key to the 'Stirps. Diop-
sidinae', Rondani gave the indication 'Thoracis aculei
quator: scutelli duo' [thorax with four spines: scutel-
lum with two] . Notwithstanding the shortness of the
genus definition, it sufficed to characterize the genus.
With 'Thoracis aculei quator' Rondani referred to the
pair of metapleural spines present in all Diopsidae
and the pair of supra-alar spines only present in
Teleopsis, while 'scutelli duo' referred to the scutellar
spines present in all Diopsidae. Some confusion was
caused by Cobiopsis latifascia (Brunetti 1928c), which
has a pair of intra-alar spines and which, due to its six
thoracic spines, was originally placed in Teleopsis (see
Feijen 1989).
Frey (1928) erected the genus Megalabops tor Diop-
sis quadriguttata Walker, though Brunetti (1928a/b)
placed this species in Teleopsis. Frey also did not pro-
duce a formal diagnosis. Only in his key to the genera
he gave for Megalabops 'Innere Orbitale nur ganz kurz
gestielt, auf der Mitte des Stiels. Die Rückendornen
ganz kurz' [IVB with a quite short base, placed in the
middle of the eye stalk. The thoracic spines quite
short]. For Teleopsis Frey stated 'Innere Orbitale auf
einem langen, dicken Stiel stehend, basal gelegen. Die
Rückendornen lang' [ivb standing on a long stalk-like
base and more basally located. The thoracic spines
short]. Steyskal (1972) placed Megalabops in synon-
ymy with Teleopsis, stating that the former only dif-
fered from the latter 'in degree of development of
same characters'. Feijen (1989) followed Steyskal's
view, stressing that for an eventual later division of
Teleopsis s. 1. different differential characters will have
to be used. Further research now revealed that sykesii,
the type-species of Teleopsis, would also fall under
Megalabops using Frey's criteria. This rather defini-
tively sinks Megalabops Frey. The confusion was
caused by '7! sykesii' Frey not being conspecific with
T. sykesii (Westwood). True Teleopsis species are so far
only recorded from the Oriental Region, while one
species is known from Madagascar. Shillito (1971a)
showed that, except for the one Madagascan species,
Teleopsis does not occur in Africa and that various
African diopsids were erroneously referred to it.
This paper should be considered as the first contri-
49
Tijdschrift voor Entomologie, volume i4i, 1998
bution towards a monograph on Teleopsis. About five
contributions are intended, in which all seventeen
recognized species will be redescribed. Besides the
two new species from Sri Lanka presently described,
thirteen additional new species from China, India,
Indonesia, Malaysia, Myanmar (Burma) and Viet-
nam will be described. This first contribution pre-
sents a genus diagnosis, a catalogue and notes on dif-
ferential and quantitative characters. In the catalogue
various new synonymies are proposed and several old
synonymies rejected. The first species treated are the
ones from Sri Lanka.
The three Teleopsis species of Sri Lanka form their
own group in the genus: the ferruginea group. A key
to the various groups recognized within Teleopsis will
be presented in a later contribution, as will be a key to
all species. Besides the three Teleopsis species, only
two other diopsids are known to occur in Sri Lanka.
Sphyracephala bipunctipennis (Senior- White 1922)
was described from Sri Lanka. It was originally placed
in Teleopsis and transferred to Sphyracephala by Feijen
(1989). The second species is a Diopsis near indica
Westwood 1837a. The Diopsis indica complex still
needs to be revised.
Abbreviations used
bmnh
BPBM
CNMS
DEIC
IVB
MCSNG
NMW
OVB
RMNH
UMO
USNM
UZMH
ZMA
British Museum (Natural History), London
Bernice P. Bishop Museum, Hawaii
National Museum Sir Marcus Fernando
Mawatha, Colombo, Sri Lanka
Deutsches Entomologisches Institut,
Eberswalde
Inner Vertical Bristle
Museo Civico de Storia Naturale 'Giacomo
Doria', Genua
Wissenschaftsbereich Zoologie, Sektion
Biowissenschaften, Martin-Luther-
Universität, Halle
Museum National d'Histoire Naturelle,
Paris
Naturhistorisches Museum, Wien
Outer Vertical Bristle
Nationaal Natuurhistorisch Museum
Naturalis, Leiden (formerly Rijksmuseum
van Natuurlijke Historie)
Hope Entomological Collections, University
Museum, Oxford
United States National Museum of Natural
History, Washington
Zoological Museum, University of Helsinki
Zoölogisch Museum, Amsterdam
Genus Teleopsis Rondani
Teleopsis Rondani (1875: 442). Type species Diopsis sykesii
Westwood 1837a, by original designation.
Megalabops Frey (1928: 70). Type species Diopsis quadrigut-
tata Walker 1856a, by original designation.
Diagnosis
Small to large diopsids with small to extremely large
eye stalks; bipartite arista; ovb varying from very short
to long, sometimes spinous; rvB varying from short to
long, base of rvB varying from small elevation to long,
stalk-like cone; facial sulcus present; facial teeth present
or absent; precoxal bridge present; supra-alar spines
present, intra-alar spines absent; scutal bristles absent;
scutellum dorsally flat or convex, scutellar spines strong
and curved, apical bristle small to medium-sized (half
the length of the spine); metapleural spines small to
medium-sized, posterolaterally directed; front femora
very slender to strongly incrassate, with two rows of tu-
bercles along most of ventral side; femora without api-
cal spurs; mid tibiae with two small subapical bristles;
alula absent, sixth vein not extending beyond anal cell,
fifth vein extending beyond posterior crossvein but not
reaching wing margin, wing varying from irrorate to
hyaline with brown bands, glabrous basal area varying
from small to large; abdomen strongly davate, slender
basal section long, syntergum consisting of terga 1 , 2
and 3, seam between terga 2 and 3 usually indistinct;
base of sternum 1 in varying degrees fused to synter-
gum, other sterna rather narrow and medially located;
spiracle 1 in tetgum or in sclerotized area between ter-
gum and sternum; between sterna 1 and 2 a small
band-like sclerite; female sterna 5, 6 and 7 usually sin-
gle sclerites (sometimes one or more are medially di-
vided); female terga 7 and sterna 7 in some species con-
nected via basal sclerotized band; female terga 8 and
sterna 8 both consisting of two sclerites; female spiracle
7 usually in membrane; male spiracle 7 often in ster-
num 7+8; three spermathecae with tooth-like orna-
ments; spermathecal ducts with U-turns near the sper-
mathecae; male sternum 6 absent; gonostyli usually
articulated, sometimes fused with periandrium; phal-
lapodeme strongly built; processus longi only running
from gonostylus to cercus (processus not distinguish-
able in species with non-articulate gonostyli); sexual
differences in eye span and front femora vary from ho-
momorphy to strong dimorphy (larger mean eye spans
and longer and more slender front femora in males).
Teleopsis is characterized as a monophyletic group
by the following groundplan conditions which are
apomorphous with respect to the gtoundplan (see
Feijen 1989) of the Diopsidae:
(1) supra- alar spines present.
(2) sternum 1 basally fused to syntergum.
(3) spiracle 1 in sclerite.
(4) spermathecal ducts with U-turns near the
spermathecae.
50
F e i j e N : Teleopsis ferrugin e a grò up
Catalogue
Teleopsis Rondani, 1875: 442
Type species Diopsis sykesii Westwood 1 837a, by
original designation.
= Megalabops Frey, 1928: 70
Type species Diopsis quadriguttata Walker
1856a, by original designation.
Ref.: Steyskal 1972: 1 1; Feijen 1989: 62.
adjacens Brunetti, 1928b: 276 (without description
in Brunetti 1928a: 272)
Holotype from Talum, Thailand (bmnh).
belzebuth Bigot, 1874: 113 {Diopsis)
Holotype from Borneo (umo).
Ref.: Bigot 1880: 94; not 'belzebuth'; Hendel
1913: 37 (= bigoti Hendel); Hendel 1914: 94;
Brunetti 1 928a: 272 (synonymy established by
E.E. Austen); Frey 1928: 72; Shillito 1971b: 301.
= discrepans Walker, 1856b: 134
In part, two of the three syntypes of discrepans
are conspecific with belzebuth. Syntypes from
Sarawak, Borneo, Malaysia (bmnh).
= fallax Bigot, 1874: 111 syn. n.
Holotype from Borneo (umo, fragment).
Ref.: Bigot 1880: 94.
longiscopium Rondani, 1875: 444
Holotype from Sarawak, Borneo, Malaysia
(mcsng).
Ref.: Brunetti 1928a: 272 (as synonym of
discrepans, a synonymy established by E. E.
Austen); Guiglia 1957: 197.
bigoti Hendel, 1914:94
Type-series from Chip-Chip, Taiwan (nmw and
bmnh).
Ref.: Hendel 1913: 37 (as belzebuth); Brunetti
1928a: 270 (junior synonym of quadriguttata)
and 272 (junior synonym of discrepans);
Matsumura 1931: 371 (as Sphyracephala
quadriguttata); Shiraki 1932: 24; Hennig 1941a:
58 (as ' Megalabops quadriguttata'); Hennig
1941c: 135 (as Megalabops bigoti); Chen 1949: 2
(as possible synonym of discrepans); Steyskal
1972: 12 (junior synonym of quadriguttata).
boettcheri Frey, 1928: 75
Type-series from Catbalogan and Panaon on
Samar and from Dansalan, Mumungan, Butuan
and Surigao on Mindanao, Philippines (uzmh).
Ref.: Tenorio 1969: 483.
breviscopium Rondani, 1875: 443
Holotype from Sarawak, Borneo, Malaysia
(mcsng).
Ref.: Brunetti 1928a: 272 (as synonym of
discrepans, a synonymy established by E.E.
Austen); Guiglia 1957: 197.
discrepans Walker, 1856b: 134 {Diopsis)
Mixed type-series from Sarawak, Borneo,
Malaysia (bmnh). Two of the three types are
conspecific with belzebuth.
Ref.: Brunetti 1907: 165; Brunetti 1928a: 270;
Chen, 1949: 2 (as possible synonym of bigoti);
Tan 1965 (in part); not Burkhardt & de la
Motte 1985: 204; not Feijen 1989.
ferruginea Roder, 1893: 235 {Diopsis)
Type from Southern Sri Lanka (mluh).
Ref.: Frey 1928: 70 (as possible Megalabops); not
'ferruginea' Curran 1936: 2 (= Cyrtodiopsis
currani, Shillito 1940: 159); Shillito 1940: 157
(as Megalabops).
krombeini sp. n.
Type-series from Sri Lanka. Holotype and
paratypes in usnm, paratypes in bmnh, rmnh
and CNMS.
maculata sp. n.
Type-series from Sri Lanka. Holotype and
paratype in bmnh, paratype in usnm.
motatrix Osten Sacken, 1882: 236
Type-series (four out of five fragments) from the
Philippines (deic).
Ref.: Wulp 1897: 193 (as possible synonym of
'sykesii'Wulp); Frey 1928: 71; Tenorio 1969; 483.
onopyxus Séguy, 1949: 67
Holotype and two paratypes from Madagascar
(mnhnp).
pharao F rey, 1928: 74
Type-series from Banahao on Luzon, Philippines
(uzmh).
quadriguttata Walker, 1856a: 37 {Diopsis)
Type-series from Malacca, Malaysia (bmnh and
mnhnp).
Ref.: Wulp 1897: 196 (as possible senior
synonym of rubicundd); Brunetti 1907: 165;
Brunetti 1 928a: 270 (in part, not Taiwan
specimens); Frey 1928: 70 (as type-species of
Megalabops); not 'Sphyracephala quadriguttata'
Matsumura 1931: 371 (= bigoti); not
'Megalabops quadriguttata 'Hennig 194la: 58 (=
bigoti); Tan 1965: 25; Steyskal 1972: 12 (as
senior synonym of bigoti); Burkhardt & de la
Motte 1985: 204.
[= fenestrata Bigot, nomen nudum. In part.
Type-series from India, umo.
Ref.: Brunetti 1907: 165; Steyskal 1972: 11.]
rubicunda Wulp, 1897: 196 (as possible synonym of
quadriguttata)
Type-series of Sukabumi, Java and Hili Madjeja,
Nias (near Sumatra) (one fragment of Nias type
in zma).
Ref.: Meijere 1908: 117; Meijere 1916b: 90;
Meijere 1924b: 204; Brunetti 1928a: 270 (Sri
51
Tijdschrift voor Entomologie, volume ui, 1998
Lanka material is krombeini); Frey 1928: 75
(doubtful identification); Frey 1934: 335; nee
Tan 1965: 27 (= sexguttata).
selecta Osten Sacken, 1882: 236
Holotype from Philippines (deic).
Ref.: Frey 1928: 77.
sexguttata Brunetti, 1928b: 275 (without description
in Brunetti 1928a: 272)
Type-series from Bukit Besar, Thailand (bmnh).
shillitoiT enoiio, 1969: 483
Type series from Sulu, Mindanao, Philippines
(holotype and paratypes in bpbm, paratypes in
USNM and bmnh).
sykesii Westwood, 1837a: 310 {Diopsis)
Type series from Hurreechunderghur, Deccan,
India (umo).
Ref.: Rondani 1875: 443; not ' sykesii'Wulp
1897: 193; Brunetti 1907: 165; Brunetti 1928a:
270 (not Borneo reference); not 'sykesii 'Meijere
1911: 366; not ' sykesii' Meijere 1916b; 89; not
'sykesii' Meijere 1919: 32; not 'sykesii 'Frey
1928: 72; not 'sykesii'Tenorio 1969: 483.
= fulviventris Bigot, 1880: 94 syn. n.
Holotype from India (umo).
trichophora Meijere, 1916b: 89 (as trichophoras)
Type-series from Sibolga and Fort de Koek,
Sumatra (lectotype and paralectotypes in zma,
paralectotypes in rmnh).
Ref.: Meijere 1916a: 40 (as trichophorus, without
description); Meijere 1916c: 132 (change in
trichophora); Meijere 1924a: 60.
Discussion
Former catalogues of Teleopsis were given by Bru-
netti (1907, Oriental), Brunetti (1928a), Descamps
(1957, Afrotropical), Steyskal (1972), Steyskal (1975,
Oriental) and Cogan and Shillito (1981, Afrotropical).
Quite a number of species were erroneously placed in,
or transferred to Teleopsis. Westwood (1837b) showed
in his figures of Diopsis wiedemanni, Diopsis erythro-
cephala and Diopsis arabica supra-alar spines. There-
fore Brunetti (1928a) and Descamps (1957) placed
the three species in Teleopsis. However, all three spe-
cies are clear Diopsis. Hennig (1941b) corrected the
error for arabica, Séguy (1955) for erythrocephala and
Shillito (1971a) for wiedemanni. Teleopsis nitida
Adams, 1903 was placed in Diopsinaby Feijen (1978)
and Cogan and Shillito (1981). Brunetti (1928a)
placed Diopsis leucochira Bezzi, 1908 and Diopsis sul-
cifrons Bezzi, 1908 by mistake in Teleopsis. Brunetti
(1928b) described truncataas a Teleopsis, but Shillito
(1940) correctly placed it in Cyrtodiopsis, though er-
roneously in synonymy with C. dalmanni (Wiede-
mann) (see Feijen 1981).
Table 1. Sexual dimorphy or homomorphy for eye span and shape offrant femur for the 19 recognized Teleopsis species. For
each species a maximum of 20 specimens was measured per sex.
Teleopsis
Ratio
Eye spar
\l body length
Ratio length/
width
No. of!
lies
of front femur
examined
5
3
Category
?
3
Category
?
3
belzebuth
0.98
2.00
4.8
6.7
10
23
motatrix
0.95
1.77
strong
4.4
57
strong
4
11
shillitoi
0.89
1.55
dimorphy
4.5
5.8
dimorphy
18
10
trichophora
0.93
1.61
5.2
6.2
12
32
15
adjacens
0.88
1.11
4.4
4.8
12
breviscopium
0.98
1.41
4.6
5.2
moderate
6
2
discrepans
1.00
1.49
moderate
4.6
5.2
dimorphy
23
39
pharao
0.86
1.34
dimorphy
4.6
5.1
3
9
rubicunda
0.82
1.25
4.0
4.9
22
14
sykesii
0.94
1.19
4.1
4.2
114
67
onopyxus
0.91
-
-
3.6
-
3
0
boettcheri
0.84
0.98
weak
4.0
4.2
16
30
ferruginea
0.84
0.98
dimorphy
4.5
4.5
homo-
16
30
krombeini
0.81
1.00
3.8
3.8
3.8
3.8
morphy
9
81
9
bigoti
0.63
0.67
91
quadriguttata
0.72
0.69
homo-
3.0
3.2
4
10
sexguttata
0.71
0.73
morphy
3.3
3.3
14
5
maculata
-
0.63
-
-
4.9
0
2
selecta
-
0.84
1
-
4.4
0
2
52
Feijen: Teleopsis ferruginea group
Descamps (1957) placed sulcifions again in Diopsis
and Shillito (1971a) and Steyskal (1972) referred leu-
cochira again to Diopsis. Descamps (1957) placed Di-
opsis dubia Bigot, 1874 by mistake in Teleopsis, but
Shillito (1971a) and Steyskal (1972) referred it to Di-
asemopsis. Senior- White (1922) described Teleopsis
bipunctipennis from Sri Lanka, but Steyskal (1975)
referred it to Pseudodiopsis and subsequently Feijen
(1989) placed it in Sphyracephala. Brunetti (1928c)
described Teleopsis latifascia from Sierra Leone, but
Shillito (1971a) rejected its placing in this genus.
Feijen (1989) erected Cobiopsis for latifascia, a mono-
typical genus in the Diasemopsis group. In the umo
collection two undescribed species of Bigot are pre-
sent: ' 'Diasemopsis fenestrata (as fenestrates) and 'Di-
asemopsis rufithorax'. The latter species represents a
Cyrtodiopsis, but the 'type-series' of fenestrata consists
of two Teleopsis and one Cyrtodiopsis. Brunetti (1907)
and Steyskal (1975) placed ' fenestrata 'nomen nudum
in synonymy with Teleopsis quadriguttata, which was
incorrect for the Cyrtodiopsis specimen, while the two
Teleopsis might belong to a species of the quadrigutta-
ta-complex.
Diopsis neesii Westwood 1837b (origin 'Japoniâ?')
could, judging from Westwood's description and fig-
ure, only have been a Cyrtodiopsis or a Teleopsis. As
Westwood clearly mentioned the presence of two pairs
of thoracic spines, the former possibility seems more
likely. The type of D. neesii kept in the Bonn Zoolog-
ical Museum was lost during the second world war.
Notes on some characters
Eye stalks (table 1). - In Teleopsis both dimorphic
(males with relatively larger eye stalks) and homo-
morphic (average relative eye span in males and fe-
males about equal) species occur (Burkhardt & de la
Motte 1985). Strongly dimorphic species in this re-
spect are T. belzebuth, motatrix, shillitoi and tricho-
phora, while adjacens, breviscopium, discrepans, pha-
rao, rubicunda, sykesii and probably onopyxus are
moderately dimorphic species. Weakly dimorphic
species are T. boettcheri, ferruginea and krombeini,
while bigoti, quadriguttata, sexguttata and probably
maculata and selecta are homomorphic species. In
table 1 data are presented on the ratio eye span/body
length and the ratio length/width of the front femur
for the 19 Teleopsis. A clear correlation appears be-
tween the degree of sexual dimorphy for these two
characters. A more detailed analysis of this phenome-
non will be presented in the last contribution to the
Teleopsis monograph, when thirteen more species will
have been described. The following categories are
used for description of the eye span: very small for a
ratio eye span/body length (e/b) 3=0.75, small in case
of0.75<E/B3=0.90, medium-sized for 0.90<^/b^ I. \0,
large for 1.10<e/b^1.25, very large for 1.25<e/b2;
1 .50 and extremely large for 1.50<e/b.
Base of ivb. - A stalk-like base (cone) of the ivb oc-
curs in T. boettcheri, breviscopium, discrepans, motatrix,
pharao, rubicunda, selecta and shillitoi. In some species
only a tiny bristle stands on this cone, but in rubicun-
da in which the cone is not very large, the bristle is well
developed. A wart-like base or just an elevation of the
stalk at the base occurs in T. adjacens, bigoti, belzebuth,
ferruginea, krombeini, maculata, onopyxus, quadrigut-
tata, sexguttata, sykesii and trichophora.
Facial teeth. - Rounded facial corners are present
in T. adjacens, bigoti, ferruginea, krombeini, maculata,
quadriguttata, rubicunda, selecta, shillitoi and tri-
chophora. Facial teeth are present in T. belzebuth, bre-
viscopium, boettcheri (very small) , discrepans, motatrix,
onopyxus, pharao, sexguttata and sykesii.
Face. — In species like T. sexguttata and quadrigut-
tata the upper half of the face is strongly protruding,
which, together with the facial sulcus, divides the face
into four sections. On the other hand, in T. ferrug-
inea frons and face are flat and smoothly rounded.
Colour. - Most species are reddish (ferrugineous)
or dark brown to blackish. As some variation occurs
in nature or due to conservation methods, care has to
be taken with this character. The legs are usually red-
dish to brown. Dark species (blackish head and tho-
rax) are T. bigoti, discrepans, onopyxus, sexguttata, shil-
litoi, and sykesii. Species with a dark thorax and red to
brown head are T. adjacens, motatrix, pharao, quadri-
guttata, selecta, shillitoi and trichophora. Reddish to
brown head and thorax occur in T. boettcheri, belze-
buth and rubicunda. The easiest species to recognize
by its colours is T. ferruginea, with its glossy black
head, reddish legs, thorax and basal abdomen and
black apical abdomen. The abdomen has a few lateral
pollinose spots or transverse bands, which have to be
examined under various angles before being described.
Wing. - The wing pattern appears a useful charac-
ter for dividing Teleopsis in groups, but care has to be
taken as some intraspecific variation might occur,
while some patterns are sometimes quite indistinct.
In most species the apex of the wing is infuscated, but
in T. selecta and sykesii a distinctly dark apex occurs,
while in boettcheri and motatrix the apex is clearly
hyaline. In the two species with a dark apex T. selecta
is easily recognized by its wholly black wing with hya-
line base and six spots, while sykesii has a very dark
preapical band, a weaker band in the middle and
some brown around the anal cell. In T. ferruginea the
whole wing is slightly infuscated with a dark preapical
band and two hyaline spots at the wing margins prox-
imally of the preapical band. In T. discrepans, mota-
trix and shillitoi there is a distinct preapical band, a
vague narrow band in the middle (which sometimes
reaches the costa) and a spot near the anal cell. In T.
belzebuth a preapical band and a band in the middle
53
Tijdschrift voor Entomologie, volume hi, 1998
Fig. 1-5. Teleopsis ferruginea, Kandy, Udawattakele Sanct. -1,9 habitus in dorsal view; 2, 8 head in anterior view; 3, â
wing; 4, 9 front leg, outer side; 5, dorsal view of 9 terga 8 & 10 and cerei; fig. 1-4 scales 1 mm, fig. 5 scale 0.1 mm.
occur which are centrally connected (H-shape). In T.
bigoti, krombeini sp. n. and (most?) rubicunda the
same H- configuration occurs, but in addition a nar-
row basal band occurs. In T. adjacens, boettcheri, tri-
chophora and (some?) rubicunda also an H-shape oc-
curs but in the cross band a third hyaline spot occurs
distally of the anterior crossvein. In T. quadriguttata
three transverse bands occur, interconnected central-
ly so that four hyaline spots result (double H-shape)
In T. sexguttata and pharao also a double H-shape oc-
curs, but with, in addition, a small hyaline spot in the
first posterior cell (if the pale apex is also included in
the count of spots the six-spotted sexguttata is ob-
tained). In T. maculata sp. n. from Sri Lanka a dark
apex, a dark preapical band and two small dark spots
in the middle of the wing occur.
Syntergum. - An unusual feature of Teleopsis is,
that in all species the base of sternum 1 is fused to the
syntergum. The degree of fusion varies. In T. ferrug-
inea, for instance, the fusion includes the anterior half
of the sternum, while externally no seam is visible. In
other species a distinct seam is present.
Hairiness. - Most Teleopsis are rather bald, with
only a few scattered white hairs. Only T. trichophora
and boettcheri have a more dense covering with large
hairs, but these species are not as hairy as Cyrtodiopsis
dalmanni (Wiedemann).
Front femora (table 1). - Intrageneric variation oc-
curs in the ratio of length/width. It varies from very
slender in the male of T. belzebuth, slender in the
males of motatrix, shillitoi, trichophora, breviscopium,
discrepans and pharao, moderately incrassate in males
of adjacens, rubicunda, sykesii, boettcheri, ferruginea,
maculata and selecta, to incrassate in males of krom-
beini, bigoti, quadriguttata and sexguttata. The follow-
ing categories are used for description of the femur:
strongly incrassate if the ratio length/width (l/V) is
2=3.0, incrassate in case of 3.0<l/w"3=4.0, moderately
incrassate for 4.0 <l/w^ 5.0, slender for 5.0<l/w^6.5,
and very slender for 6.5<l/w. In a number of species
sexual dimorphy occurs in the shape of the front
femora. In T. belzebuth, motatrix, shillitoi and tricho-
phora the females have more incrassate front femora
and this can be described as strong dimorphy. This
strong dimorphy in the shape of the front femora is
correlated to the strong dimorphy in the eye span
54
Feijen: Teleopsis ferruginea group
(table 1). In T. adjacens, breviscopium, discrepant,
pharao, rubicunda, sykesii and probably onopyxus the
femora in the females are only somewhat more in-
crassate as in the males and this can be referred to as
moderate dimorphy. In T. boettcheri, ferruginea,
krombeini, bigoti, quadriguttata, sexguttata and proba-
bly maculata and selecta no differences between the
sexes occur in the shape of the front femora.
Key to the diopsids of Sri Lanka
1. Arista tripartite; alula present; sixth vein extend-
ing beyond anal cell; syntergum only including
terga 1-2, apical bristles several times longer
than scutellar spines
Sphyracephala bipunctipennis
- Arista bipartite; alula absent; sixth vein not ex-
tending beyond anal cell; syntergum including
terga 1+2+3, apical bristles smaller than one-
fourth the length of the scutellar spine or absent.
(Diopsinae) 2
2. Supra-alar spines absent, scutellar spine almost
straight, apical bristle absent, wing with only an
apical wingspot Diopsis near indica
- Supra- alar spines present, scutellar spines strong-
ly curved, apical bristles smaller than one- fourth
the length of the scutellar spine, wing pattern
otherwise ( Teleopsis) 3
3. Central head glossy black, only one, preapical,
wing band, front femora with dark spots 4
- Central head glossy brown, upper half of face
more pronounced, thorax brown with posterior
half of scutum glossy, three crossbands on the
wing, front femora not with spots, dorsal ab-
domen yellowish brown basally and dark brown
more apically, tergum 3 with two pale spots
basally Teleopsis krombeini sip. n.
4. Frons and face flat and smoothly rounded, thorax
(including collar) reddish, one very broad preapi-
cal crossband on the wing, dorsal abdomen
brown with dark brown semicircular spot on apex
Teleopsis ferruginea
- Upper half of face more pronounced, collar
glossy black, scutum chestnut brown with black
sutures, wing with two small dark spots in the
middle and distinct apical spot, one narrow V-
shaped preapical crossband on the wing, ab-
domen blackish brown with posterolateral pale
spots on the first five terga
Teleopsis maculata sp. n.
The ferruginea group
All three known species originate from Sri Lanka.
The group is especially characterized by the small ar-
ticulate gonostyli without microtrichia but with hairs.
Furthermore, the ferruginea group is characterized by
the absence of facial teeth, minute to small ivb, small
base of ivb, medium-sized ove-, female spiracle 7 in
membrane, divided male sternum 5, large and broad
male cerei, weak sexual dimorphy in eye span and sex-
ual homomorphy in front femora. The other groups
presently recognized {motatrix group, rubicunda
group, sykesii group, boettcheri group, quadriguttata
group and sexguttata group) will later be character-
ized. Intrageneric phylogeny will be discussed in the
final contribution to the monograph.
Teleopsis ferruginea (Roder, 1 893)
(figs. 1-11)
Diopsis ferruginea Roder, 1893: 235. Holotype 9 South Sri
Lanka (Ceylon), v. 1889, H. Fruhstorfer (mluh). [Exam-
ined]
ÌMegalabops ferruginea; Frey, 1928: 70.
Megalabops ferruginea; Shillito, 1940:157.
Teleopsis ferruginea, Steyskal, 1972: 11.
[Misidentification. - Teleopsis ferruginea; Curran, 1936: 2
(= Cyrtodiopsis currani Shillito, 1940)].
Further material. - 49, 6â , Kandy, Udawattakele
Sanct., Kan. Dist., Sri Lanka, 6-8.vi.1978, K V. Krombein,
P. B. Karunaratne, T. Wijesinhe, V. Kulasekare, L. Jaya-
wickrema; 29, 3d, Udawattakele Sanct., Kan. Dist., Sri
Lanka, 8-1 l.ii. 1979, K. V. Krombein, P. B. Karunaratne, T.
Wijesinhe, S. Siriwardane, T. Gunawardane; 19, 3d,
Kandy Reservoir Jungle, Kan. Dist., Sri Lanka, 10. ii. 1979,
K. V. Krombein, P. B. Karunaratne, T. Wijesinhe, S. Siri-
wardane, T. Gunawardane; 39, 6d, 1?, Udawattakele
Sanct., Kan. Dist., Sri Lanka, l-3.ix.1980, K.V. Krombein,
P.B. Karunaratne, T. Wijesinhe, L. Jayawickrema, V. Gu-
nawardane (all Krombein material in usnm, except for some
specimens in cnms and rmnh); 1 9 , 1 d , Sri Lanka (bmnh);
5d, Sri Lanka, Dr. Thwaites, 67-25 (bmnh); 29, 2d, 3?,
Sri Lanka, Weston Coll., bmnh 1924-199; 19, 2d, Per-
adeniya, Sri Lanka, 30.iv.1891, Lt Col. Yerbury (bmnh);
2d, Henaratgoda, Sri Lanka, i. 1901 (bmnh); 1?, Sudugan-
ga, 10.ix.l919,R. Senior White (bmnh). In total 169, 30d
and 5? were examined.
Description
Measurements. - length of body in 9 5.5 mm ± se
0.2 (range 4.3-6.2) and in cT 5.8 mm ±0.1 (range
4.7-6.2), eye span in 9 4.6 mm ± 0.2 (range 3.8-5.2)
and in o* 5.9 mm ± 0.2 (range 4.0-7.4), length of
wing in 9 4.3 mm ±0.1 (range 3.4-4.8) and in 6 4.4
mm ±0.1 (range 3.8- 5.1), length of scutellar spine in
9 1.16 mm ± 0.03 (range 0.99-1.30) and in 6 1.16
mm ± 0.02 (range 0.90-1.30).
Head. - Central part glossy black (figs. 1-2); ocellar
tubercle glossy black; frons very smooth with laterally
at base of stalk a deep groove (fig. 2); arcuate groove
narrow and indistinct; face flat and smooth, laterally
and ventrally with a typical 'woolly' type of pollinosi-
ty, some small white hairs, facial sulcus indistinct, fa-
cial corners rounded; mouthparts greyish brown; eye
55
Tijdschrift voor Entomologie, volume i4i, i99S
Fig. 6-1 1. Teleopsis ferruginea, Kandy, Udawattakele Sanct. - 6, ventral view of $ postabdomen; 7, ventral view of subanal
plate; 8, spermathecae; 9, posterior view of periandrium with gonostyli and cerei; 10, lateral view of phallapodeme and aedea-
gus; 11, ejaculatory apodeme and sac; fig. 6 scale 1 mm, fig. 7-11 scales 0.1 mm.
span small in female (eye span 16% smaller than
length of body) and medium-sized in male (eye span
2% larger than length of body), stalks yellowish
brown, anteriorly and posteriorly with a blackish
band, apices blackish pollinose; ivb minute (almost
invisible) and spinous, base of ivb a small elevation,
one quarter the diameter of the eye stalk; ovb medi-
um-sized, 2x the diameter of the eye stalk, spinous.
Thorax. - Uniformly reddish brown (ferrugi-
neo us), anterior edge of collar black, laterally two
black stripes on collar, prosternum with medial black
spot between front legs, scutellar spines darker apical-
ly; dorsally thinly pollinose, anterior edge of collar
densely pollinose, scutellar spines also pollinose but
supra- alar spines glossy, upper half of pleura thinly
pollinose, lower half glossy except for pollinose poste-
rior area, sterna glossy but thinly pollinose anteriorly
and near legs; supra-alar spines medium-sized, almost
3x as long as metapleural spines, laterally directed,
somewhat turned upward; scutellar spines medium-
sized, almost 3x scutellum, moderately curving up-
ward and outward (fig. 1), diverging under an angle
of 80°; metapleural spines blunt, posterolaterally di-
rected; apical bristle small, about one-sixth of length
of scutellar spine; few small hairs on thorax, the hairs
on the scutellar spines without basal warts.
Wing. - apex almost hyaline; very broad, very dark
preapical band (fig. 3), apically convex, proximally
concave and slightly extending in the first posterior
cell; remainder of wing slightly infuscated but base
hyaline, also hyaline spots in marginal and submar-
ginal cell and basally in second posterior cell, both
these hyaline spots adjoining the preapical band;
glabrous basal area includes basal and posterior parts
of costal cell, small basal section of marginal cell, basal
half of first basal cell, most of second basal cell and
posterior half of anal cell.
Legs. - Coxa 1 brown with blackish bands on inner
and outer side, femur 1 brown with small dark spot
basally on inner side, large black spots on distal two-
thirds of outer side and some black dorsally (fig. 4),
tibia 1 black, tarsi 1 yellowish white; mid leg brown
with dark bands on femur and darker base of tibia;
hind leg dark brown; femur 1 moderately incrassate
56
Feijen: Teleopsis ferruginea group
in 9 and 3, ratio of length/width in 9 4.5 ± 0.0
(range 4.4-4.8) and in 3 also 4.5 ± 0.0 (range 4.2-
5.0), tubercles on distal three-quarters, inner row in
female with 25.7 tubercles ± se 0.6 (range 22-30)
and in male with 27.6 tubercles ± 0.4 (range 24-31),
outer row in female with 21.9 tubercles ± 0.4 (range
19-25) and in male with 22.6 tubercles ± 0.4 (range
19-26); legs covered with some hairs, ventral side of
front femur densely covered with small hairs.
Preabdomen. - Syntergum dorsally yellowish
brown basally, apex with semi-circular dark brown
spot (fig. 1), tergum 4 dark brown except for lateral
edges proximally, other terga dark, dark areas of terga
together forming a circle which coincides with the
dark preapical wingbands (with wings in folded posi-
tion); two basal terga thinly pollinose except for two
pairs of glossy lateral spots, tergum 3 basally with two
large pollinose spots laterally; dark apical terga polli-
nose; sterna yellowish brown, sternum 1 glossy; seam
between terga 2 and 3 indistinct; basal half of ster-
num 1 smoothly fused to syntergum (fig. 6), inter-
sternite 1-2 very short, other sterna located medially;
spiracle 1 in sclerite (fig. 6).
Female postabdomen. - Strongly deflexed; terga 6
and 7 single rectangular sclerites (fig. 6); tergum 8
consisting of two sclerites (fig. 5), sclerites anteriorly
glabrous; tergum 10 ill-defined, with one pair of
hairs; cerei rather elongate, ratio of length/width 3.6,
covered with microtrichia and with a number of
hairs; sternum 6 a single rectangular sclerite; sternum
7 posteriorly constricted medially, giving two sclerites
joined anteriorly; spiracle 7 in membrane; sternum 8
represented by two triangular sclerites; subanal plate
(fig. 7) pentagonal with rounded corners, at apex a
pair of large hairs; laterally some small hairs, covered
with microtrichia; spermathecae (fig. 8) rounded
with few rounded protuberances, ten in the single
theca and eight each in the pair, heavily sclerotized;
genital ring tapering towards one side.
Male postabdomen. - Sternum 4 a single rectangu-
lar sclerite; sternum 5 consisting of two small sclerites;
sternum 7+8 without sclerotized connection to anteri-
or sclerites of periandrium; spiracle 7 in sternum 7+8;
periandrium (fig. 9) rounded, with about 14 pairs of
hairs, covered with microtrichia; gonostyli articulated,
remarkably small, apically rounded, a few hairs, no
microtrichia, gonostyli connected to lateral side of
cerei, not interconnected; cerei large, broad, flat,
somewhat rectangular with rounded corners, ratio of
length/width 1.9, covered with microtrichia and hairs;
phallapodeme (fig. 10) with broad anterior arm,
abruptly narrowing anteriorly and strongly curving
downward, anterior arm slightly longer than posterior
arm; aedeagus (fig. 10) with rather long ejaculatory
duct sticking out from apex; ejaculatory apodeme fan-
shaped (fig. 11), ejaculatory sac relatively small. The
interpretation of the ejaculatory duct as sticking out
from the apex needs confirmation, as Kotrba (1993)
states for Cyrtodiopsis that the ejaculatory duct opens
in a phallotrema at the base of a long process.
Diagnosis
Teleopsis ferruginea gives its name to the ferruginea
group. It is the most colourful of all diopsids with its
glossy black head, reddish legs, reddish thorax and
basal abdomen and black apical abdomen. Further-
more it can be recognized by its wing pattern (one
broad preapical crossband), distribution of microtri-
chia on the wing (small glabrous area), minute ivb,
medium-sized ovb, small base of ivb, absence of facial
teeth, moderately incrassate front femora, rectangular
female sternum 6, almost completely divided female
sternum 7, female spiracle 7 in membrane, rounded
pentagonal subanal plate, rather elongate female cer-
ei, round spermathecae with 8-10 rounded protuber-
ances, articulated and very small gonostyli without
microtrichia and with few hairs, large and broad male
cerei, anteriorly curved phallapodeme with longer an-
terior arm, fan-shaped ejaculatory apodeme, weak
sexual dimorphy in eye span and sexual homomorphy
in front femora.
Habitat
The two Kandy locations are jungle areas at an alti-
tude of 550-650 m and have an average annual rain-
fall of 1950 mm.
Teleopsis krombeini sp. n.
(figs. 12-23)
Type material. - 3 holotype, 49 , 66 and 2? para-
types from Thawalamtenne, Kan. Dist., 2200 ft, Sri
Lanka, 4.ix.l980, K. V. Krombein, P. B. Karunarat-
ne, T. Wyesinhe, L. Jayawickrema, V. Gunawardane
(usnm). 1 9 paratype from Kitugala, Bandarakele Jun-
gle, Keg. Dist., Sri Lanka, 17-18.iii.1979, K. V.
Krombein, P. B. Karunaratne, T. Wyesinhe, L. Jaya-
wickrema, V. Gunawardane (usnm); 3 9 paratypes,
Kandy, Sri Lanka, 28.V.1892, Lt Col. Yerbury (two
with rubicunda label, det. J. A. Tenorio) (bmnh); 2c?
paratypes, Haragam, Sri Lanka, 24. v. 1892, Lt Col.
Yerbury (bmnh); 1 3 (bmnh genitalia slide), Kandy,
Sri Lanka, 24.V.1892, Lt Col. Yerbury; 19 (bmnh
genitalia slide), Haragam, Sri Lanka, 1. vi. 1892, Lt
Col. Yerbury (both slides were identified as rubicunda
by J. Smart). Some paratypes are deposited in rmnh
and CNMS. In total 9 9 , 9cT and 2? were examined.
Description
Measurements. — Length of body in 9 5.7 mm ±
SE 0.2 (range 5.2-6.2) and in cT 5.8 mm ± 0.2 (range
4.9-6.4), eye span in 9 4.6 mm ± 0.1 (range 4.0-5.1)
57
Tijdschrift voor Entomologie, volume ui, 1998
Fig. 12-16. Teleopsis krombeini — 12, ? paratype, Kitugala, habitus in dorsal view; 13, 3 holotype, Thawalamtenne, head in
anterior view; 14-15, â paratype, Thawalamtenne; 14, wing; 15, front leg, outer side; 16, $ paratype, Thawalamtenne, lat-
eral view of abdomen; fig. 12-15 scales 1 mm, fig. 16 scale 0.1 mm.
and in â 5.8 mm ± 0.4 (range 4.2-6.8), length of
wing in ? 4.2 mm ±0.1 (range 3.8-4.6) and in â 4.3
mm ±0.1 (range 3.6- 4.7), length of scutellar spine in
9 1.16 mm ± 0.04 (range 1.02-1.30) and in S 1.12
mm ± 0.04 (range 0.90-1.24).
Head. - Central part glossy brown, concolorous
with stalks (fig. 12), laterally some pollinosity; frons
smooth, slightly concave, with laterally at base of stalk
a groove (fig. 13); arcuate groove dark brown, broad,
area along arcuate groove wirh small transverse
grooves; upper half of face more pronounced, face
with ridge parallel to and just below arcuate groove,
centrally two small protuberances, facial corners
rounded; eye span small in female (eye span 19%
smaller than length of body) and medium-sized in
male (eye span just as long as length of body), stalks
glossy brown, apices black pollinose; ivb rather small,
spinous, equal in size to diameter of eye stalk, ivb on
small tubercle with a length of less than half the di-
ameter of the stalk; ovb medium-sized, slightly longer
than ivb, spinous.
Thorax. - Uniformly brown; collar dorsally glossy,
laterally pollinose; scutum thinly pollinose dorsally,
but posterior half glossy; scutellum pollinose, scutel-
lar spines dark, glossy; pleura glossy but pollinose
near base of wing and posteriorly; sterna glossy with
some pollinosity between front legs; supra-alar spines
medium-sized, twice as long as metapleural spines,
laterally and somewhat upward directed; scutellar
spines long, more than 3.5x scutellum, strongly curv-
ing upward and outward (fig. 12), diverging under an
angle of 90°; metapleural spines blunt, posterolateral-
ly directed; apical bristle small, about one-fifth of
length of scutellar spine, medially directed (lacking in
most specimens); few, small hairs on thorax, hairs on
scutellar spines without basal warts.
Wing. - Apex infuscated; three crossbands, broad
preapical dark band and broad central dark band
joined in first posterior cell, forming H-configuration
(fig. 14), anterior part of preapical band and part in
first posterior cell darker, some parts of central band
darker; narrow, vaguely infuscated, basal band run-
ning from tip of costal cell to tip of anal cell, region
below anal cell infuscated with darker central
smudge; the pattern of bands leaves a hyaline base,
adjoining hyaline spots in submarginal, marginal and
discal cell, a hyaline spot centrally in the third poste-
rior cell, a hyaline spot in submarginal and marginal
58
Feijen: Teleopsis ferruginea group
Fig. 17-23. Teleopsis krombeini paratypes, Thawalamtenne -17, ventral view of 9 postabdomen; 18, ventral view of subanal
plate; 19, spermathecae; 20, S sternum 7+8; 21, posterior view of periandrium with gonostyli and cerei; 22, lateral view of
phallapodeme and aedeagus; 23, ejaculatory apodeme and sac; fig. 17 scale 1 mm, fig. 20 scale 0.2 mm, all others scales 0. 1 mm.
cell and a hyaline spot basally in second posterior cell;
the glabrous basal area includes the costal cell, base of
submarginal cell, basal halves of first and second basal
cell and almost the whole anal cell; the adjoining hya-
line spots in submarginal, marginal and discal cells are
also glabrous.
Legs. - Coxa 1 brown, femur 1 brown with darker
inner side, tibia 1 blackish brown, tarsi 1 brown; mid
and hind legs brown, apical quarter of femora darker,
basal half of mid tibia and whole of hind tibia black-
ish brown; femur 1 incrassate (fig. 15) in both sexes,
ratio of length/width 3.8 ± se 0.1 in ? (range 3.4-
4.2) and 3.9 ± 0.1 in cT (range 3.6-4.1), tubercles on
distal three-quarters, inner row in female with 26.8
tubercles ± se 0.7 (range 23-30) and in male with
28.0 tubercles ± 0.6 (range 23-31), outer row in fe-
male with 23.4 tubercles ± 0.5 (range 20-26) and in
male with 24.9 tubercles ± 0.4 (range 22-27); legs
covered with some hairs.
Preabdomen. - Tergum 1 and 2 glossy yellowish
brown with two transverse darker bands in the mid-
dle; tergum 3 and subsequent terga dark brown,
glossy (some very thin pollinosity) , tergum 3 with
two pale spots basally, covered with white pollinosity,
white pollinosity extending laterally (fig. 12); sterna
yellowish brown, sternum 1 glossy; seam between ter-
ga 2 and 3 not very distinct; basal half of sternum 1
smoothly fused to syntergum (fig. 17), intersternite
1-2 narrow, other sterna located medially; spiracle 1
in sclerite (fig. 17).
Female postabdomen. — Strongly deflexed; terga 6
and 7 single rectangular scleri tes (fig. 17), tergum 7
concave posteriorly; tergum 8 represented by two scle-
rites (fig. 16), sclerites anteriorly glabrous; tergum 10
with one pair of hairs; cerei very elongate, ratio of
length/width 5.4, covered with microtrichia and with
a number of hairs; sterna 5 and 6 rectangular sclerites;
sternum 7 more or less U-shaped, the central-posteri-
or region not being sclerotized; spiracle 7 in mem-
brane; sternum 8 represented by two small sclerites;
subanal plate (fig. 18) in between pentagonal and
heart-shaped, with posteriorly two pairs of long hairs
and about five pairs of short hairs, covered with mi-
crotrichia; spermathecae (fig. 19) rounded with sub-
59
Tijdschrift voor Entomologie, volume ui, 1998
Fig. 24-29. Teleopsis maculata $ holotype, Nuwara Eliya - 24, habitus in dorsal view; 25, head in anterior view; 26, wing
27, front leg, outer side; 28, front femur, inner side; 29, lateral view of abdomen; scales 1 mm.
apically a few rounded protuberances (four in the sin-
gle theca and two each in the pair), heavily sclerotized.
Male postabdomen. - Sternum 4 a single rectangu-
lar sclerite; sternum 5 represented by two square scle-
rites; sternum 7+8 without sclerotized connection to
anterior sclerites of periandrium; left spiracle 7 in
sternum 7+8, fight spiracle 8 in lateral slit of sternum
7+8 (fig. 20); periandrium (fig. 21) rounded, with
about 1 3 pairs of rather short hairs, covered with mi-
crotrichia; gonostyli small, articulated, apically some-
what pointed, hairs mainly on inner side, no mi-
cro trichia, gonostyli connected to lateral side of cerei,
not interconnected; cerei large, rather broad, flat,
somewhat rectangular with rounded corners, ratio of
length/width 2.2, covered with microtrichia and
hairs, medial-posterior corners with narrow, darkly
sclerotized, region; phallapodeme (fig. 22) with
broad, rather straight anterior arm about equal in
length to posterior arm, vane broad; aedeagus (fig.
22) with rather long ejaculatory duct sticking out
from apex; ejaculatory apodeme fan-shaped (fig. 23),
ejaculatory sac relatively small.
Diagnosis
Teleopsis krombeini belongs to the ferruginea group
and can be recognized by its wing pattern (three cross-
bands), distribution of microtrichia on the wing
(glabrous basal area and basal anterior spot), small
IVB, medium-sized ovb, small base of ivb, absence of
facial teeth, incrassate front femora, glossy posterior
half of scutum, rectangular female sternum 6, U-
shaped female sternum 7, female spiracle 7 in mem-
brane, pentagonal to heart-shaped subanal plate, very
elongate female cerei, round spermathecae with 2-4
protuberances, small and articulated gonostyli with-
out microtrichia, large and broad male cerei, phal-
lapodeme with straight anterior arm equal in length
to posterior arm, fan-shaped ejaculatory apodeme,
weak sexual dimorphy in eye span and homomorphy
in front femora.
Habitat
The Thawalamtenne location is a jungle area at an
altitude of about 600 m and with an average annual
rainfall of 1750 mm.
60
Feijen: Teleopsis ferruginea group
Fig. 30-33. Teleopsis maculata â paratype, Hakgala Natural Reserve - 30, ventral view of basal abdomen; 31, posterior view
of periandrium with gonostyli and cerei; 32, lateral view of phallapodeme and aedeagus; 33, ejaculatory apodeme and sac; fig.
30 scale 1 mm, fig. 31-33 scales 0.1 mm.
Teleopsis maculata sp. n.
(figs. 24-33)
Type material. - 6 holotype, Nuwara Eliya, Sri
Lanka, l4.vii.1892, Lt Col. Yerbury, (bmnh); 3
paratype from Hakgala Natural Reserve, N.E. Dist.,
Sri Lanka, 6-7.ii.1979, K. V. Krombein, P. B.
Karunaratne, T. Wijesinhe, S. Siriwardane, T. Gu-
nawardane (usnm); 1? paratype (no head and ab-
domen), Punda luoya (Pundaloya), Sri Lanka, E. E.
Green, bmnh 90- 1 1 5.
Description
Measurements. - Length of body in â 5.8 mm
(range 5.6-5.9), eye span 3.7 mm (range 3.4-3.9),
length of wing 4.7 mm (range 4.2-5.2), length of
scutellar spine 1.14 mm (range 1.12-1.15).
Head. - Central part glossy black (figs. 24-25); frons
with medial depression below ocellar tubercle, bor-
dered by two elevated areas, frons laterally surrounded
by circular groove, two slightly paler spots above arcu-
ate groove (fig. 25), anterolaterally roughened areas; ar-
cuate groove dark, rather deep and narrow with rough-
ened surface; face glossy black, dorsally around facial
sulcus fine grooves, upper half more pronounced, fa-
cial corners rounded; eye span very small in male (eye
span 37% smaller than length of body), stalks yellow-
ish brown, dark stripes anteriorly on ventral part,
broad apices black pollinose; rvB small, just longer than
diameter of eye stalk, on vague elevation of about one
quarter the length of the diameter of the stalk; ovb
medium-sized, Wix. the length of the rvB, spinous.
Thorax. - Collar glossy black, laterally some polli-
nosity; scutum chestnut brown with blackish sutures,
posteriorly darker, thinly pollinose; scutellum brown,
thinly pollinose, scutellar spines darker, glossy; meso-
pleuron and sternopleuron blackish brown anteriorly,
remainder of pleura brown, near abdomen and in su-
tures blackish, pleura pollinose but most of sterno-
pleuron, central part of mesopleuron and central part
of hypopleuron glossy; sterna glossy but pollinose an-
teriorly and posteriorly; supra-alar spines medium-
sized, slightly more than twice the length of the meta-
pleural spine, laterally directed, slightly turned upward;
scutellar spines long, almost 3.5x scutellum, rather
straight, pointing upward and outward (fig. 24), di-
verging under an angle of 80°; metapleural spines
posterolateral^ directed; apical bristle small, about
one-fourth of length of scutellar spine, slightly turned
inward; few, small hairs on thorax, small hairs along
scutellar spines on tiny warts.
Wing. - Distinct apical wingspot in submarginal
and first and second posterior cell (fig. 26); dark
preapical band, somewhat V-shaped; tiny dark spots
in the middle of the marginal cell and around the an-
terior crossvein; remainder of wing lightly infuscated
with two pale spots proximally of the preapical band
near the wing margins; the small glabrous basal area
61
Tijdschrift voor Entomologie, volume ui, 1998
includes most of the costal cell, base of submarginal
cell, basal halves of first and second basal cell and
basal half of anal cell.
Legs. - Coxa 1 , trochanter 1 and femur 1 yellowish,
femur 1 (figs. 27-28) with two brown spots on inner
side, tibia 1 and metatarsus 1 dark brown, remainder
of tarsus 1 paler (fig. 27); mid leg yellowish brown, fe-
mur 2 with dark spots on inner and outer side in the
middle and dark apical third, basal half of tibia 2
slightly darker; hind leg yellowish brown, femur 3
with long dark spots on inner and outer side in the
middle and dark apical quarter, tibia 3 mainly dark
brown with pale brown band in the middle; femur 1
moderately incrassate in â , ratio of length/ width 4.9
± 0.0, range 4.9-5.0, tubercles on distal three-quarters,
inner row in S with 28.0 ± 0.4 tubercles (range 27-
29), outer row with 25.0 ± 0.4 tubercles (range 24-26).
Preabdomen. - Terga blackish brown with pos-
terolaterally pale spots on terga 1, 2, 3, 4 and 5 (figs.
24, 29), pollinose except for glossy basal half of ter-
gum 2 and glossy basal band of tergum 3, pale spots
on tergum 3 covered with dense pollinosity; sterna
yellowish brown, sternum 1 glossy; seam between ter-
ga 2 and 3 almost invisible; basal edge of sternum 1
linked to syn tergum (fig. 30), intersternite 1-2 nar-
row, other sterna located medially; spiracle 1 in ter-
gum (fig. 30).
Male postabdomen. - Tergum 6 with anterior con-
cavity medially; sternum 4 a single rectangular sclerite;
sternum 5 represented by two sclerites; sternum 7+8
without sclerotized connection to anterior sclerites of
periandrium; left spiracle 7 in sternum 7+8, right spir-
acle 8 just in membrane; periandrium (fig. 31) round-
ed, with about 1 5 pairs of hairs, covered with mi-
crotrichia; gonostyli small, articulated, apically blunt,
hairs distributed along the sides, especially long hairs
on inner side, no microtrichia, gonostyli connected to
lateral side of cerei, not interconnected; cerei medium-
sized, broad, flat, rectangular, ratio of length/width
2.0, covered with microtrichia and hairs, apical hairs
long, medial-posterior corners with darkly sclerotized
area; phallapodeme (fig. 32) anteriorly curved down-
ward, anterior arm broad and somewhat longer than
posterior arm, aedeagus (fig. 32) with rather long ejac-
ulatory duct sticking out from apex; ejaculatory
apodeme fan-shaped (fig. 33), ejaculatory sac small.
Diagnosis
Teleopsis maculata belongs to the ferruginea group
and can be recognized by its wing pattern (apical spot
and one V-shaped crossband), distribution of micro-
trichia on the wing (small glabrous area), small ivb,
medium-sized ovb, small base of ivb, absence of facial
teeth, glossy black central head and collar, moderate-
ly incrassate front femora with two spots on inner
side, abdomen with five pairs of lateral spots, small
and articulated gonostyli without microtrichia and
with long hairs on inner side, broad male cerei, ante-
riorly curved phallapodeme with anterior arm slightly
longer than posterior arm, fan-shaped ejaculatory
apodeme and probably minor sexual dimorphy in eye
span and sexual homomorphy in front femora.
Habitat
The average annual rainfall at Nuwara Eliya, sever-
al miles away from the montane rain forest on Mount
Hakgala, is 2 1 60 mm.
Acknowledgements
I am grateful to the following curators for the op-
portunity offered to study the Teleopsis in their collec-
tions: A. Albrecht (uzmh), B. Brugge (zma), R. Con-
treras-Lichtenberg (nmw), M. Dorn (mluh), N. E.
Evenhuis (bpbm), P. Grootaert (irsnb), P. J. van Hels-
dingen (rmnh), K. V. Krombein (usnm), L. Matile
(mnhnp), G. C. McGavin (umo), B. Pitkin (bmnh)
and V. Raineri (mcsng). D. Burkhardt and I. de la
Motte (University of Regensburg) placed material
from Malaya and Sarawak at my disposal.
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Received: 18 August 1997
Accepted: 13 January 1998
63
Tijdschrift voor Entomologie, volume i4i, 1998
continuation from page 48
many species can be identified. Unfortunately the key
in both the German and French edition is the same as
in the previous edition, with the German species only
and therfore cannot be used in more southern re-
gions. The German version can be used to the north-
ern border of the Mediterranean, including eastern
France, northern Italy, Slovenia and northern Croat-
ia. The French edition contains most continental
French species, but again lacks several of the more
eastern species, included in the German edition. So to
be 'complete', it is necessary to buy both versions.
The 'new' photographs in the French edition are not
as good as the original ones, but it contains a lot of ex-
tra information on the French Fauna. Both guides
contain brief descriptions of sound and some oscillo-
grams.
The Dutch book [now almost sold out] is not a
field-guide, it is too heavy, but more a distribution at-
las with extensive information on all 45 Dutch species
and some information on 1 9 species from neighbour-
ing countries or exotic introduced species. Apart from
the Dutch maps, the book contains European distrib-
ution maps, graphs on phenology and biotopes, and
descriptions of sound with oscillograms. Also all
species are illustrated with an artists impression of the
species and a colour photograph. It contains also an
illustrated key of about 55 western European species.
The three cd's all have a smaller number of species
than those of Ragge & Reynolds, but those of Bonnet
and Odé have a much better sound quality because
they are based on digital recordings. Bonnet's record-
ings have a nice sound, because they often are made
in nature, and also background sounds can be heard
such as those from birds, frogs, and other orthopter-
ans. The cd by Odé has relatively long fragments,
and is the only one with a considerable amount of
other sound elements that the calling songs. The Ger-
man cd is a simple transmission of the earlier musi-
cassette and treats the German species only.
In short all these cd's have their own right, which
also holds for the books. For identification the Bell-
mann guides are indispensible, but for southern Eu-
rope still incomplete. With the Ragge & Reynolds
cd's at least some of this area is covered. The real Or-
thopterist probably would like to have all these books,
which make this interesting group one of the better
known groups in northwestern Europe. Now it is
waiting for field guides or identification books for
southern parts of Europe, highly needed to replace
the outdated books by Harz. I hope that there are
some orthopterists who dare to take this challenge.
[Erik J. van Nieukerken]
64
SlGFRlD INGRISCH
Bad Karlshafen, Germany
A REVIEW OF THE ELIMAEINI OF WESTERN
INDONESIA, MALAY PENINSULA AND THAILAND
(ENSIFERA, TETTIGONIIDAE, PHANEROPTERINAE)
Ingrisch, S., 1998. A review of the Elimaeini of Western Indonesia, Malay Peninsula and Thai-
land (Ensifera, Tettigoniidae, Phaneropterinae). - Tijdschrift voor Entomologie 141: 65-108,
figs. 1-174, maps 1-3.[issn 0040-7496]. Published 30 November 1998.
A review of the Elimaeini occurring in Western Indonesia, Malay Peninsula, Thailand and adja-
cent islands is given. The stridulatory file on the underside of the male left tegmen, conchate
phallus sclerites and modifications of the gonangulum of the ovipositor are introduced as new di-
agnostic characters. The phylogenetic relations of the genera and subgenera are discussed.
Hemielimaea Brunner, 1878 is reduced to a subgenus of Elimaea Stài, 1874. Phaneroptera
pammpunctata Serville, 1839 is designated as type species of Rhaebelimaea. In the subgenus Eli-
maea, there are two distinct species groups differing by the width of the tegmen which is (a)
wider than the pronotal length {Elimaea s. str.) or (b) narrower than the pronotal length in males
and of subequal width in females (E. poaefolia-gioup) . A tabular key to the species in the area
considered is provided; the diagnostic characters are figured. Nine species are described as new:
Elimaea (Rhaebelimaea) mentaweii sp. n., E. (R.). maninjauensis sp. n., E. (R.). modiglian i i sp. n.,
E. (R.). pseudochloris sp. n., E. (R.). sinuatasp. n., E. (R.). apicata sp. n., E. (R.). pentaspina sp. n.,
Elimaea (Elimaea) thaii sp. n., E. (Elimaea) nautica sp. n.. Two species of the E. poaefolia-group
which are only known from single females, are informally described but not named. Elimaea mi-
nor (Brunner, 1891), formerly included in E. (Rhaebelimaea), is transferred to E. (Orthelimaea),
comb, n., E. (R.) curvicercata (Brunner, 1891) is removed from synonymy under E. (R.) pamm-
punctata (Serville, 1839). Elimaea subcarinata (Stil, 1861) is removed from synonymy under E.
chloris (De Haan, 1 842) and regarded a valid species with E. appendiculata Brunner, 1 878 as a
new synonym (the latter is not identical with E. punctifera (Walker, 1869)). E. chloris and espe-
cially E. punctifera have a much more restricted distribution than previously thought. The female
oi E. (Hemielimaea) procera (Ingrisch, 1990), and the male of E. (H.) cuculiata (Ingrisch, 1990)
are described for the first time. Information on stridulation of Ectadia fidva Brunner, 1893, Eli-
maea (E.) chloris, E. (E.) subcarinata, E. (E.) thaii and E. (H.) cuculiata is given.
S. Ingrisch, Eichendorrfweg 4, D-34385 Bad Karlshafen. E-mail: sigfrid.ingrisch@real-net.de
Key words. — Elimaeini, Phaneropterinae, South East Asia, phylogeny, key to species, taxono-
my, new species, stridulation
The Phaneropterinae were divided into several
groups by Brunner (1878) and these may be regarded
as tribes in present day taxonomy (e.g. Bei-Bienko
1965). The tribe Elimaeini Yakobson, 1905 (group
Elimaeae Brunner, 1878) consists so far of only three
genera, Elimaea Stal, 1874, Hemielimaea Brunner,
1878, and Ectadia Brunner, 1878, all occurring in the
Oriental region. It is thought to be most closely relat-
ed with the African and Mediterranean Acrometopini
and the Oriental Mirolliini (Brunner 1878).
The genus most rich in species, Elimaea Stai, 1874,
contains a few widespread and numerous local species
(Brunner 1878, 1891, Kirby 1906, Hebard 1922a,
Karny 1926a-c, 1931, Tinkham 1943, Bei-Bienko
1951, 1962, 1965, Ingrisch 1990a, Jin & Xia 1994).
It was divided into three subgenera by Karny (1926a):
Orthelimaea, Rhaebelimaea, and Elimaea s. str. While
the anterior femur is straight in Orthelimaea, it is
curved in Elimaea and Rhaebelimaea. The latter sub-
genera differ in the radius sector branching before or
about in the middle of the tegmen. Hemielimaea dif-
fers from Elimaea by the tibial tympanum which is
open on external and covered by a conchate fold on
internal side, while in Elimaea it is covered by a con-
chate fold of the integument on both sides. Hemieli-
maea species are so far only known from China and
Indochina, while the distribution of Elimaea s. lat.
reaches from India to the Sunda Islands and the
Philippines (map 1).
The species of the Elimaeini belong to the com-
65
Tijdschrift voor Entomologie, volume i4i, 1998
mon Phaneropterinae in South East Asia. They can
be found in primary forests as well as in secondary
vegetation. They live in the lower vegetation as shrubs
and grasses and are thus amongst the first katydids to
be met when looking for Orthoptera in tropical Asia.
Their activity is largely nocturnal. During day time
they are often found sitting along the central vein of a
leaf with the fore legs and antennae stretched anteri-
orly. In doing so, the head fits in the phasmid-like
curvature of the anterior femora which is found in
some of the taxa. The middle and the hind legs are
pressed together and spread in an acute angle from
the body (figs. 152-153, 155).
The great species diversity of Elimaea in South East
Asia was noted by Hebard (1922a) and Karny
(1926a). Despite this, many species are not well de-
scribed, and it is difficult to identify Elimaea species
without re-examining the types or specimens from
the type localities. This is especially true for the
species of the subgenus Elimaea s. str. which cannot
be identified with certainty on the basis of the previ-
ously used characters. Thus it became necessary to
find new differentiating characters. The characters
that were most commonly used to identify Elimaea
species are the tegminal venation (i.e. the branching
of the radius sector), male and female subgenital
plates, male cerei, and coloration. Coloration can,
however, be greatly variable within species. The sub-
genital plates also show some individual variation
(compare e.g. figs. 57 and 58) and, unfortunately, its
shape in museum specimens may have changed due to
desiccation after death (compare e.g. figs. 59 and 60).
Examination of material from the museums in Bo-
gor (Indonesia), Leiden (The Netherlands) and Ge-
noa (Italy) and my collection from Thailand, Sumatra
and Java revealed two facts: (1) The number of species
and the regional diversity is even greater than previ-
ously thought; and (2) there are additional distinctive
characters that allow - together with those previously
described - a clear distinction between the species in
both sexes. Those characters are, in males: (1) the
stridulatory file on the underside of the left tegmen,
and (2) sclerotised structures of the phallus which
consist of a pair of conchate (mussel-shaped) sclerites
with serrate margin and sometimes with an addition-
al unpaired sclerotised projection. Those sclerotised
structures are characteristic for the species of Rhaebe-
limaea and Hemielimaea. In females, the gonangulum
(a lateral sclerite at the base of the ovipositor) can be
provided with an appendage of variable shape at the
ventral margin. In some cases the corresponding area
of the ventral ovipositor valves is also modified.
The purpose of the present paper is to show the
distinctiveness of the new and previously recognised
characters. This may be the basis for a future revision
of the genus. The results are presented in form of a
tabular key to the species from Java, Sumatra, Menta-
wei Islands, Malay Peninsula and Thailand. The re-
striction to that area is for practical reasons, as no new
material from other areas has been available to me.
Especially the species from Vietnam and China can-
not be revised without extensive new material as sev-
eral of the types are lost (see checklist below). The
present paper includes most of the species previously
assigned to Rhaebelimaea, except those from Borneo
and the Philippines, and it includes a redescription of
the types of the "classical" species of Elimaea s. str.
which were most difficult to differentiate and often
wrongly identified.
The consideration of the new diagnostic characters
makes it necessary to discuss the relations between the
genera and subgenera anew on a phylogenetic basis. A
checklist of the taxa of the Elimaeini is provided. De-
scriptions of species are restricted to new or insuffi-
ciently known species.
Depositories
Material examined for this study is deposited in the
following collections:
BMNH The Natural History Museum, London,
England
ci Private collection of S. Ingrisch, Bad Karls-
hafen, Germany, will later be deposited in a
museum
dab Department of Agriculture, Entomology
and Zoology Division, Chatuchak, Bangkok,
Thailand
MCSN Museo Civico di Storia Naturale, Genoa, Italy
MZB Museum Zoologicum Bogoriense, Bogor,
Indonesia
NHMW Naturhistorisches Museum, Vienna, Austria
MNHN Museum National d'Histoire naturelle, Paris,
France
RMNH Nationaal Natuurhistorisch Museum, Leiden,
The Netherlands
NRS Naturhistoriska Riksmuseet, Stockholm,
Sweden
SMF Forschungsinstitut und Naturmuseum
Senckenberg, Frankfurt/M, Germany
zsi Zoological Survey of India, Calcutta, India.
Methods for recording stridulation
Stridulation of one species each of Ectadia, Hemieli-
maea and Rhaebelimaea and that of three morphologi-
cally similar species of Elimaea s. str. were so far record-
ed. Recording of the song of E. chloris was done in the
field, that of the other species in wooden cages with
gauze walls in the laboratory. In Phaneropterinae, there
is usually a male-female response stridulation (Heller
1990). With one exception, only the male songs which
are more distinctive were studied. All recordings were
66
s
OC 1*3
t
©
El
©
A
Ingrisch: Review of Elimaeini
<3
es
,;2
^
^
<
<U
•^
-a
O
=<
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m
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ai
Fig. 1. Three hypothetical phylogenetic trees (A-C) of the Elimaeini without Ectadia as discussed in the text. Circles = apomorphic
characters; squares = plesiomorphic characters; inverse numbers = apomorphic character must have evolved twice independently.
Characters; 1, tegmen with parallel principle veins and strong, regular, vertical transverse veinlets; 2, gonangulum of ovipos-
itor ventrally projecting or with appendage; 3, anterior femur straight; 4, tibial tympana conchate on both sides; 5, tibial tym-
pana conchate on internal, open on external side; 6, phallus with a pair of conchate sclerites; 7, anterior femur curved;
8, tegmen very elongate (narrower than pronotum length in male); 9, phallus sclerites secondarily reduced; 10, radius sector
branching in anterior quarter or third of tegmen.
made during the night in darkness. For recording in
the field, a portable cassette recorder (Sony WM3) with
a stereo microphone (Sennheiser mke 66) and Cr02-
tape cassettes were used (frequency range 0.05 - 15
kHz), in the laboratory a cassette recorder (Kenwood
KX 880 hx) with a mono microphone (akg D202 or
Sennheiser black fire 541) and metal tape cassettes (fre-
quency range 0.05 - 20 kHz). Analysis of stridulation
was done using the programme Soundscope on a Mac-
intosh PC (Quadra 840) with a MacAdios 11/16 board
as a/d converter which was connected via a MacAdios
ABO box to the cassette recorder. The sound was fil-
tered before analysis (zoiA-filter wlfdap). The oscillo-
grams were copied into a graphic program for arrange-
ment and final presentation.
Phylogeny
Within the Elimaeini, Ectadia is rather remote
from the other genera and shows some similarities
with the Acrometopini and the Mirolliini. The simi-
larities of Ectadia with Acrometopa concern e.g. the
rather stout and somewhat breakable antennae (but
not as extreme as in Acrometopa) and the wing dimor-
phism of male and female (the wings of Ectadia fe-
males are however less reduced than those of Acrome-
topa females). The tegminal venation with an irregu-
lar course of the media and a wide network of oblique
transverse veins resembles the situation in Mirollia. In
the other Elimaeini, the antennae are thin and flexi-
ble, the tegmina of both sexes are of equal size, and
the principle veins of the tegmen are straight, subpar-
allel and rather equally spaced from each other with
the transverse veins running vertical to the principle
veins. The relations of Ectadia have to be re-evaluated
in a suprageneric revision of the Phaneropterinae,
which is beyond the scope of the present paper. Brun-
ner (1891) assigned also the African genus Pantolepta
Karsch, 1889 to the Elimaeini, because it has the fore
and mid femora compressed and spined on ventral
margins. It was said to be close to both Acrometopa
and Ectadia Brunner (1891). The stout, breakable an-
tennae and the reduced hind wings of the female,
which do not surpass the tegmina, makes it more like-
ly that Pantolepta belongs to the Acrometopini, but
its position should also be re-evaluated in a supra-
generic revision. The remaining taxa of the Elimaeini
are certainly closely related to each other, and these
should be considered, if the formal, present day divi-
sion into genera and subgenera is correct, from a phy-
logenetic point of view.
A phylogenetic tree based on the presently recog-
67
Tijdschrift voor Entomologie, volume i4i, 1998
nised division is presented in fig. 1 a. The armature of
the tibial tympana is obviously a character that is
readily modified in the Phaneropterinae. It is con-
diate on both sides in the Acrometopini, in Ectadia
and in Elimaea, and it is conchate on internal and
open on external side in Hemielimaea and in the
Mirolliini. As such there is not a very strong argu-
ment for a separation of Hemielimaea and Elimaea.
Moreover, if we agree with the traditional arrange-
ment of the genera, we have to suppose that the oc-
currence of a pair of conchate sclerites on the phallus
has evolved twice independently in this closely related
group of taxa (in Hemielimaea and in Rh aeb elimaea).
The phallus of the majority of Phaneropterinae is ba-
sically simply membranous. Sclerotised structures ex-
ist in several genera but they are not well known.
Their structures probably evolved independently in
different genera or genus groups. However, those of
Hemielimaea and Rhaebelimaea species are very simi-
lar in shape and position and may thus be regarded as
homologous structures of both taxa. In Orthelimaea,
the phallus is membranous, but in some species the
dorsal lobes are provided with a stiffened longitudinal
ridge covered with spinules. A situation which is re-
garded more primitive than the possession of distinct
sclerites as in Hemielimaea and Rhaebelimaea. Sclero-
tised structures are also found on the phallus of the
Mirolliini, but these are quite different from those of
the Elimaeini. They will be dealt with in a later paper.
If however, we regard Hemielimaea and Rhaebeli-
maea as sister groups on the base of the phallus scle-
rites as a synapomorphic character (fig. lb), we have
to suppose that the phasmid-like curvature of the an-
terior femora evolved twice independently in a close-
ly related group of taxa {Rhaebelimaea and Elimaea).
Curved anterior femora are unique in the Phanero-
pterinae and may thus be regarded as a synapomor-
phic character of Rhaebelimaea and Elimaea.
A solution would be a sister group relationship as
presented in the third phylogenetic tree (fig. lc). In
this tree, both characters, phallus sclerites and curved
anterior femora, are considered to be apomorphic and
evolved only once within Elimaeini. But in contrast
to the first tree (fig. la), it is supposed that the phal-
lus sclerites are secondarily reduced in Elimaea s. str.
The third tree is the most parsimonious, as we do not
have to suppose that any obviously homologous char-
acters evolved twice independently.
If we accept the phylogenetic tree in fig. 1 c to be
correct, the division of the Elimaeini into genera and
subgenera has to be changed. There are two alterna-
tives: (1) At least Orthelimaea has to be removed from
Elimaea and raised to full generic rank, as it is more
remote from Elimaea and Rhaebelimaea than is
Hemielimaea. Rhaebelimaea might also be raised from
a subgenus under Elimaea to full generic rank as it
shows close relations to both, Hemielimaea (conchate
phallus sclerites) and Elimaea (curved anterior femo-
ra). (2) Hemielimaea should become a subgenus of
Elimaea. As this is not a complete revision of the
tribe, the second alternative is preferred.
The species previously included in the subgenus
Elimaea can be divided into two distinct morpholog-
ical groups, as already recognised by Karny (1926a):
(1) species with moderately narrow tegmina which
are distinctly wider than the pronotum length, and
(2) species with very elongate tegmina which are in
the male narrower than the pronotum length and in
the female of subequal width. To the former belongs
the type species {Phaneroptera subcarinata Stài, 1861)
of Elimaea and they are thus the Elimaea s. str. The
latter are listed here as the E. poaefolia-group within
the subgenus Elimaea. As both are sister groups with
many shared characters, they can remain in the same
subgenus. If the males, which are unknown for sever-
al species of the E. poaefolia- group, show constant
differences to Elimaea s. str., it might later be useful
to erect a separate subgenus for this group.
Systematic part
Checklist of the species of Elimaeini
With information on types, type localities and de-
positories. Abbreviations: H holotype, S Syntypes, F
female, M male; synonyms marked by =.
Ectadia Brunner, 1878
fulva Brunner, 1 893
hm: Burma: Carin Cheba (Genoa)
= abbreviata Brunner, 1 893
hf: Burma: Carin Ghecu (Genoa), synonymy by Bei-Bi-
enko (1954)
pilosa Brunner, 1878
hm: Kashmir (Vienna)
sulcataXiz&C Liu, 1990
hm: Yunnan: Baoshan (Shanghai)
Elimaea { Orthelimaea) Karny, 1 926
flavolineata Brunner, 1 878
hm: Sri Lanka (Vienna)
bimalayana Ingrisch, 1990b
hm: Nepal: Prov. Narayani, Bhainse Dobhan (Munich)
hunanensis Kang & Yang, 1 992
hf: China: Hunan (Beijing)
insignis (Walker, 1 869) {Phaneroptera)
hm: Bangladesh: Silhet (London)
= notabilis (Walker, 1 869) {Phaneroptera)
hf: Bangladesh: Silhet (London), synonymy by Kirby
(1906) and Uvarov (1927)
= annulata Brunner, 1878
sfm: "Hinter-Indien", Calcutta (Vienna; Budapest, lost),
synonymy by Kirby (1906) and Uvarov (1927)
inversa Brunner, 1891
hm: Sulawesi (Warsaw? formerly coll. Dohrn, Stettin)
68
Ingrisch: Review of Elimaeini
klinghardti Krausze, 1 903c
hm: Tonkin (lost)
leeuwenii Kamy, 1926a
SMM: Thailand: Bangkok (Bogor; Leiden)
/»/'«or Brunner, 1891
smm: Java: Tengger mountains (Vienna)
securigera Brunner, 1 878
Sfm: East-India: Simla (Geneva)
Elimaea (Hemielimaea) Brunner, 1893
cbinensis (Brunner, 1 878) {Hemielimaea)
sfm: China (Budapest, lost?; Vienna)
cuculiata (Ingrisch, 1990a) (Hemielimaea)
hf: Thailand: Kanchanaburi prov., Erawan falls (Frank-
furt/M)
formosana (Shiraki, 1930) (Hemielimaea)
sfm: Taiwan: Arisan Mt. (Ent. Mus. Govern. Res. Inst.,
Taiwan)
mannhardti Krausze, 1 903a
hf: Tonkin: Than-Moi (lost), combined with Hemieli-
maea by Dohrn 1 906
procera (Ingrisch, 1990a) (Hemielimaea)
hm: Thailand: Chanthaburi prov., Khao Soi Dao
(Frankfurt/M)
tonkinensis (Dohrn, 1906) (Hemielimaea)
sfm: Tonkin: Than-Moi (Warsaw? formerly Stettin)
Elimaea (Rhaebelimaea) Karny, 1926
adspersa Dohrn, 1906 (Elimaea signata vzr. adspersa)
sfm: Sumatra: not further specified, but Dohrn collected
in North Sumatra (Warsaw? formerly Stettin)
apicata sp. n.
hm: Thailand: Surat Thani province, Khao Sok
(Bangkok)
bakeri Hebard, 1922a
hm: Mindanao: Davao (Philadelphia)
bidentata Brunner, 1878
hf: Malabar (Berlin)
brunneri Dohrn, 1906
hf: Philippines (Vienna), replacement name for E.
parumpunctata Brunner, 1878 nee Serville, 1839
caricifolia (De Haan, 1842) (Phaneroptera)
hm: Borneo: Loetontoer (Leiden)
= femorata Brunner, 1878
hf: Borneo (Vienna), synonymy by Dohrn (1906) and
Karny (1923)
curvicercata Brunner, 1891
sfm: Java orientalis (Vienna)
filicauda Hebard, 1922a
hm: Luzon: Laguna, Los Banos (Philadelphia)
hebardi Karny, 1 926b
sfm: South Sumatra: Lampongs, Wai Lima (Bogor;
Leiden)
kraussi Karny, 1 926a
sfmm: West Java, Cibodas, 1400m (Bogor; Leiden)
lamellipes Hebard, 1922a
hf: Sabah: Labuan (Philadelphia)
longicercata Brunner, 1891
sfm: Borneo (Vienna)
malayica Karny, 1920
sfm: Borneo (Vienna), replacement name for E. poaefo-
lia Brunner, 1 878 nee De Haan, 1 842
maninjauensis sp. n.
hm: West Sumatra: Maninjau (Bogor)
mentaweii sp. n.
hm: Mentawei Islands (Bogor)
modiglianii sp. n.
hm: Sumatra: Si-Rambe (Genoa)
moultonii Karny, 1923
hf: Sarawak: Baram river (Singapore?)
neglecta Karny, 1 926c
hf: Malaysia: Selangor, Kanching (Kuala Lumpur)
parumpunctata (Serville, 1839) (Phaneroptera)
hf: Java (Paris)
pentaspina sp. n.
HF: Thailand: Chanthaburi prov., Khao Soi Dao
(Bangkok)
pseudochloris sp. n.
hm: Thailand: Trang prov., Khao Chong (Bangkok)
puncticosta Bolivar, 1914
hf: ? Philippines or Himalaya (lost), occurs in the
Philippines according to Karny (1926b)
roseoalata Brunner, 1 89 1
hf: Sumatra: Deli (Warsaw? formerly coll. Dohrn, Stettin)
siamensis Karny, 1 926c (Elimaea signata siamensis)
hf: South Thailand: Nakhon Sri Tamarat (Kuala Lumpur)
signata Brunner, 1 878
sfm: Singapore: Bukit (Vienna)
sinuata sp. n.
hm: Mentawei Islands: Sipora (Bogor)
spinigera Brunner, 1 878
hm: Singapore: Bukit (Vienna)
sumatrana Karny, 1 926a
hm: West Sumatra: Padangische Bovenland, Batu
Sangkar (Philadelphia), replacement name for E.
parumpunctata Hebard, 1922a nee Serville, 1839
transversa Ingrisch, 1990a
hf: Thailand: Chanthaburi prov., Khao Soi Dao (Frank-
furt/M)
willemsei Karny, 1 926b
hm: South Sumatra: Lampongs, Wai Lima (Leiden)
The subgeneric position of the following species
was not considered before. Judging from the descrip-
tions, they might belong to Rhaebelimaea:
melanocantha (Walker, 1 869) (Phaneroptera)
hm: Sri Lanka (London)
= cannata Brunner, 1 878
hf: Sri Lanka (Berlin), synonymy by Uvarov (1927)
nigrosignata Bolivar, 1 900
sffmm: South India: Madure, Kodaikanal (Madrid, Paris)
verrucosa Brunner, 1 878
sfm: ? (Geneva)
Elimaea (Elimaea) Stal, 1874
annamensis Hebard, 1 922a
hm: Annam: Phuc-Son (Philadelphia)
berezovskii Bei-Bienko, 1951
hm: Sichuan: Lunnfu-Kuochikow (St. Petersburg)
chloris (De Haan, 1842) (Phaneroptera)
sfmm: Java: Thihanjavar (Leiden)
fallax Bei-Bienko, 1951
hf: Southern Maritime Territory of USSR (St. Petersburg)
hoozanensis Karny, 1915
sfm: Taiwan: Hoozan (DEI Berlin-Dahlem)
nautica sp. n.
hm: Thailand: Chanthaburi prov., Khao Soi Dao
(Bangkok)
69
Tijdschrift voor Entomologie, volume i4i, 1998
punctifera (Walker, 1869) {Phaneroptera)
hm: Bangladesh: Silhet (London)
= diversa (Walker, 1869) {Phaneroptera)
hf: Bangladesh: Silhet (London) synonymy by Uvarov
(1927)
schmidtiYLrausze, 1903a
hm: Annam: Phuc-Son (lost)
semicirculata Kang & Yang, 1992
hm: Fujiang, Dehua (Beijing)
setifera Bei-Bienko, 1962
hm: Yunnan (St. Petersburg)
subcarinata (Stài, 1861) {Phaneroptera)
sfm: Hongkong (Stockholm)
= appendiculata Brunner, 1878
hm: Indochina (Vienna) synonymy in this paper
= rubicunda Krausze, 1 903a
hm: Tonkin: Than-Moi (lost) synonymy by Dohrn
( 1 906) as synonym of E. appendiculata
thaiisp. n.
hm: Thailand, Tak prov.: Mae Salid, Monkrating, 700m
(Bangkok)
tympanalis (Matsumura & Shiraki, 1908) (Phaneroptera)
smm: Taiwan (Matsumura's coll.)
The subgeneric position of the following species
was not stated with the description nor later revised.
Most of them probably belong to Elimaea s. str., but
some might belong to other subgenera:
atrata Carl, 1914
hm: Tonkin (Geneva)
cheni Kang & Yang, 1 992
hm: China: Hunan (Beijing)
Hi Kang & Yang, 1992
hm: China: Guangxi, Longgang (Beijing)
obtusilota Kang & Yang, 1992
hm: China: Guangxi, Longzhou (Beijing)
schenklingi Karny, 1915
sfm: Taiwan, Koshun (DEI Berlin-Dahlem)
lanceolata (Walker, 1859) {Steirodon)
hf: Sri Lanka (London)
= rufonotata Walker, 1 869 nee Serville, 1 839 {Phanero-
ptera) Sri Lanka (London) synonymy by Kirby 1906
macra (Serville, 1839) {Phaneroptera)
hm: ? (Paris)
marmorata Brunner, 1878
hf: Sumatra (Vienna), included in Rhaebelimaea by
Karny (1926a), but judging from the shape of the
tegmen, it might belong in the £. poaefolia-group
nigerrima Krausze, 1903a
hf: Tonkin: Than-Moi (lost)
parva Liu, 1993
hm: China: Fujian, Mt. Longqi (Shanghai)
rufonotata (Serville, 1839) {Phaneroptera)
hm: Bombay (Paris)
terminalis Liu, 1993
hm: China: Fujian, Mt. Longqi (Shanghai)
Elimaea poaefolia-group
jacobsonii Karny, 1926a
hm: West Sumatra: Bukittinggi, Fort Kock (Leiden)
poaefolia (De Haan, 1 842)
sfm: Java (Leiden)
rosea Brunner, 1 878
sfm: Borneo (Vienna; Dresden)
sp. 1
F: Thailand: Chiang Mai prov., Chiang Dao (Bangkok)
sp. 2
F: Thailand, Tak prov., Mae Salid, Monkrating, 700m (ci)
Species with uncertain status:
aliena (Walker, 1 869) {Phaneroptera)
hm: Bangladesh: Silhet (London), treated in Kirby
(1906) as a synonym of E. parumpunctata; this is cer-
tainly wrong; if it belongs in Elimaea, it might belong to
the poaefolia-group
theopoldi Krausze, 1903b
hf: Tonkin (lost), cited in Karny (1926a) as "species du-
bia".
triticifolia (De Haan, 1842) {Phaneroptera)
hf: Borneo: Kahayan (Leiden), listed under Elimaeam
Brunner (1878) and Kirby (1906); according to Karny
(1926a) this is not an Elimaea species but might belong
to IHabra
Tentative key
This key is meant for the species of Elimaeini oc-
curring in Java, Sumatra, Mentawei Islands, Malay
Peninsula, Singapore and Thailand.
Although the key is largely based on material studied
by myself, not all species have been seen by me. Some
illustrations were taken from Hebard (1922a) or Karny
(1926a-c) and adapted in size to the original drawings.
1. Anterior tibia with internal (= anterior) tympa-
num covered by a conchate fold, external (= pos-
terior) tympanum free. Hemielimaea 2
- Anterior tibia with internal and external tympa-
num covered by a conchate fold 3
2. Tegmen long and narrow (in S 6.7X, $ 6.2-
7. IX longer than wide). Stridulatory file with a
strong step-like declination slightly behind mid-
dle of length (figs. 14-15). Conchate sclerites of
phallus forming a quarter of a circle (fig. 95). Fe-
male subgenital plate with apex in middle slight-
ly concave (fig. 118). Eastern Central Thailand ..
E. (H.) procera
- Tegmen relatively broader (in S 5.7-6.2 X, in 9
5. 8-6. OX longer than wide). Stridulatory file
with a weaker step-like declination slightly be-
hind middle of length (figs. 16). Conchate scle-
rites of phallus forming almost a semicircle (fig.
96). Female subgenital plate with apex in middle
subtruncate to slightly convex (fig. 117). Central
Thailand E. (H.) cuculiata
3. Tegmen: radius sector fused with media; media
with zigzag course and sending several oblique,
subparallel branches to posterior margin of teg-
men. Male: tenth abdominal tergite prolonged
behind with apex subtruncate. Female: wings
shorter than in male but hind wings still slightly
surpassing tegmina. Burma, North Thailand,
South China Ectadia fulva
70
Ingrisch: Review of Elimaeini
- Tegmen: radius sector separate; media straight,
running about in middle between radius (respec-
tively radius sector) and cubitus; between radius
and cubitus with vertical, rather regularly spaced,
transverse veinlets. Male: tenth abdominal tergite
either not prolonged or if prolonged, apex divid-
ed into 2 or 3 lobes. Female: wings of same size as
in male 4
4. Anterior femur not compressed, straight (fig.
151). Orthelimaea 5
- Anterior femur in basal area compressed and
phasmid-like curved (figs. 153, 162) 6
5. Male cerei sickle-shaped, bent in basal third, af-
terwards flattened (fig. 52). Male subgenital plate
split for less than half of its length (fig. 78).
Stridulatory file with circa 30 teeth (fig. 2). Fe-
male subgenital plate triangular. Ovipositor
sabre-shaped, margins with large teeth (fig. 119).
Thailand E. (O.) leeuwenii
- Male cerei gradually curved, slightly widened be-
fore apical cone (fig. 51). Male subgenital plate
split for more than half of its length (fig. 79).
Stridulatory file with circa 70 teeth (fig. 3). Fe-
male unknown. Eastern and Central Java
E. (O.) minor
6. Phallus with a pair of conchate sclerites (figs. 85-
94). Tegmen with radius sector branching about
in or behind middle of tegmen (figs. 152-155,
159, 162), rarely (E. pseudochloris) before middle
(fig. 161). Pronotum often, but not in all species,
slightly constricted in circa middle of length 7
- Phallus membranous without sclerites. Tegmen
with radius sector branching distinctly before
middle of tegmen (figs. 156-158, 160, 164).
Pronotum not constricted in middle. Elimaea
37
7. Tegmen slightly widening towards apex (width in
middle 6 mm, at apex 7 mm). Pronotum with
lateral margins of disc angularly rounded. Suma-
tra (only known from the female type)
E. (?) marmorata
- Tegmen narrowing towards apex or of subequal
width throughout. Pronotum with lateral mar-
gins of disc rounded. Rhaebelimaea 8
8. Tegmen with radius sector branching before mid-
dle of tegmen (fig. 161). Male subgenital plate
step-like constricted behind base, thereafter nar-
row, parallel-sided (fig. 73). Stridulatory file with
large, spaced teeth throughout (fig. 9). Phallus
sclerites as in fig. 93. South Thailand
E. (R.) pseudochloris
- Tegmen with radius sector branching in or be-
hind middle of tegmen (figs. 152-155, 159, 162).
Male genitalia of different shape 9
9. Pronotum and single cells of tegmen densely suf-
fused with black dots, giving the impression of
the tegmen being marbled with somewhat irregu-
lar, quadrangular, black spots. Male cerei surpass-
ing subgenital plate, round, before apex slightly
widened. Male subgenital plate apically pro-
longed into a long, very narrow projection with
bulging lateral margins, split in circa apical quar-
ter with lobes not deviating. Sumatra
E. (R.) adspersa
- Pronotum and tegmen less densely covered with
black dots. Male external genitalia of different
shape 10
10. SS 11
- 9 9 25
1 1 . Tenth abdominal tergite largely prolonged be-
hind and apex divided into 2 or 3 lobes 12
- Tenth abdominal tergite not prolonged behind,
apex entire (subtruncate, slightly concave or con-
vex) 14
12. Tenth abdominal tergite with apical projections
long and narrow, forming a three-spined apex.
Malaysia, Singapore E. (R.) spinigera
- Tenth abdominal tergite with apical projections
bilobate, rounded 13
13. Subgenital plate split for about apical half into 2
narrow lobes. Borneo, Sumatra? [only known
from a male and a female syntype which are pos-
sibly not conspecific] E. (R.) malayica
- Subgenital plate with apical half entire, only at
very apex faintly excised (fig. 74). South Thailand
E. (R.) apicata
14. Subgenital plate bowl-shaped with a conical, nar-
row, apical part; almost split to base (fig. 71) ..15
- Subgenital plate of different shape, split in apical
quarter or at apex only 16
15. Cerci with black apical cone long and narrow
(fig. 29). South Sumatra E. (R.) willemsei
- Cerci with black apical cone shorter and stout (fig.
30). Phallus with an unpaired ventral sclerite in
addition to the paired conchate sclerites (fig. 90).
Stridulatory file sinuate with teeth spaced in circa
basal half and dense in circa apical half (fig. 5).
Mentawei Islands E. (R.) sinuata
16. Cerci long, surpassing or embracing subgenital
plate (figs. 38-41) 17
- Cerci short, usually shorter than subgenital plate
(figs. 31-35) 20
17. Cerci regularly curved in middle; apical area sub-
straight, apical cone long and narrow; apex acute
(fig. 38). Singapore, Riouw, Pulau Ubin
E. (R.) signata
- Cerci re-curved at apex or apical cone short and
stout (figs. 39-41) 18
1 8. Cerci only in apical area distinctly curved (fig. 41).
Subgenital plate with apex of apical lobes trans-
versely truncate (fig. 75). South-East Sumatra
E. (R.) hebardi
71
Tijdschrift voor Entomologie, volume i4i, 1998
- Cerci already curved from middle of length (figs.
39-40). Subgenital plate with apical lobes round-
ed (figs. 76-77) 19
19. Subgenital plate split in circa apical third; base
broad (fig. 77). Cerci at apex re-curved with mar-
gins gradually tapering (fig. 40). Stridulato ry file
with teeth spaced in basal area (fig. 7). Conchate
sclerites of phallus as in figs. 85-86. Mentawei Is-
lands E. (R.) mentaweii
- Subgenital plate split only at apex; base narrow
(fig. 76). Cerci before apex slightly widened and
then suddenly constricted to apical cone (fig. 39).
Stridulatory file with teeth rather dense from base
(fig. 8). Conchate sclerites of phallus as in fig. 94.
West Sumatra E. (R.) maninjauensis
20. Subgenital plate with apical lobes step-like con-
stricted and apex spiniform (fig. 70). North-East
Sumatra E. (R.) roseoalata
- Subgenital plate with apex of different shape... 21
21. Tegmen with dorsal field considerably widened
and on right tegmen with subparallel transverse
veins (fig. 17). Stridulatory file very long, teeth
gradually changing from spaced teeth at base to
dense teeth at apex; at apex with a few spaced teeth
(fig. 10). Conchate sclerites of phallus as in fig. 92.
Cerci acute-angularly curved with a long, narrow,
apical part (fig. 34). West Java (mountainous)
E. (R.) kraussi
- Tegmen with dorsal field not considerably
widened and on right tegmen without subparallel
transverse veins. Stridulatory file shorter. Cerci
less strongly curved (figs. 31-33, 35) 22
22. Subgenital plate with apical part narrowly divid-
ed; apex of lobes transversely truncate (figs. 66,
68). Java 23
- Subgenital plate with apical part broader, round-
ly excised (figs. 69, 72). Sumatra 24
23. Cerci with apical cone longer and more slender
(fig. 32). Stridulatory file with circa 100 teeth;
teeth larger and more spaced (fig. 6). Conchate
sclerites of phallus higher (fig. 87). West Java
E. (R.) parumpunctata
- Cerci with apical cone shorter and stouter (fig.
33). Stridulatory file with circa 80 teeth; teeth in
central area more dense (fig. 11). Conchate scle-
rites of phallus very narrow (fig. 88). East Java....
E. (R.) curvicercata
24. Subgenital plate with apical lobes longer and more
widely spaced (fig. 69). Cerci more strongly curved
(fig. 35). West Sumatra E. (R.) sumatrana
- Subgenital plate with apical lobes shorter and
closer approached to each other (fig. 72). Cerci
less strongly curved (fig. 31). Stridulatory file
step-like declined before middle (fig. 12-13).
Conchate sclerites of phallus as in fig. 89. North-
West Sumatra E. (R.) modiglianii
25- Subgenital plate with both, long baso-lateral and
long apico-lateral projections (figs. 101, 124).
Gonangulum of ovipositor with ventro-apical
projection pointing apicad (fig. 134). Eastern
Central Thailand E. (R.) pentaspina
- Subgenital plate with apico-lateral projections
only. Gonangulum of ovipositor with ventro-api-
cal projection pointing ventrad (figs. 132-133,
135-136; gonangulum not described for all
species!) 26
26. Subgenital plate constricted before apical lobes
which are pointing laterally (figs. 103-104) 27
- Subgenital plate not constricted or apical lobes
pointing apically (figs. 97-100, 102, 105-108) ....28
27. Subgenital plate at apex more narrowly excised,
apical lobes shorter (fig. 104). South-East Suma-
tra E. (R.) hebardi
- Subgenital plate at apex widely excised, apical
lobes very long (fig. 103). North-East Sumatra...
E. (R.) roseoalata
28. Subgenital plate distinctly narrowed towards
apex; apex roundly excised between apico-lateral
lobes (figs. 105-106, 108) 29
- Subgenital plate not narrowed towards apex; if
slightly narrowed widely truncate between apico-
lateral lobes (figs. 97-100, 102, 107) 31
29. Subgenital plate with apex rather deeply excised,
apical lobes spiniform (fig. 106). Malaysia: Selan-
gor E. (R.) neglecta
- Subgenital plate with apex slightly excised (figs.
105, 108) 30
30. Subgenital plate with apico-lateral lobes acute
(fig. 108). Singapore, Riouw, Pulau Ubin
E. (R.) signata
- Subgenital plate with apico-lateral lobes circa rec-
tangular (fig. 105). West Java (mountainous)
E. (R.) kraussi
31. Subgenital plate with apex almost trilobate, simi-
lar to fig. 97, but medial lobe triangular and
emarginate. East Java E. (R.) curvicercata
- Subgenital plate of different shape; if apex almost
trilobate then medial lobe obtuse, not emarginate
(fig. 97) 32
32. Ventral valves of ovipositor at base (between sub-
genital plate and gonangulum) with a tongue-
shaped projection (figs. 121, 136). Subgenital
plate as in fig. 97. West Java
E. (R.) parumpunctata
- Ventral valves of ovipositor at base without
tongue-shaped projection (figs. 122-123) 33
33. Subgenital plate largely widening apicad; apex
broad- roundly excised (fig. 107). South Thailand
E. (R.) siamensis
- Subgenital plate not or only slightly widening
apicad; apex more or less truncate between apical
lobes or triangularly excised (figs. 98, 102) 34
72
Ingrisch: Review of Elimaeini
34. Subgenital plate with apico-lateral lobes large
(figs. 99-100). Cerci sinuate, apex subacute (figs.
143-144) 35
- Subgenital plate with apico-lateral lobes small
(figs. 98, 102). Cerci simply curved, apex obtuse
(figs. 140-141) 36
35. Subgenital plate with apico-lateral lobes pointing
apico-laterally, apex transversely truncate in be-
tween (fig. 99). West Sumatra
E. (R.) maninjauensis
- Subgenital plate with apico-lateral lobes pointing
apically; apex triangularly excised in between (fig.
100). Mentawei Islands E. (R.) mentaweii
36. Subgenital plate with apex truncate between apico-
lateral lobes but slightly excised in middle (fig. 98).
Eastern Central Thailand E. (R.) transversa
- Subgenital plate with apex convexly truncate be-
tween apico-lateral lobes (fig. 102). North-West
Sumatra E. (R.) modiglianii
37. Tegmen long and narrow, its width in middle
shorter than length of pronotum (6) or of about
equal length (9); tegmen parallel-sided or in
some males slightly narrowed in middle and
widening towards apex (about 1 mm). Pronotum
with lateral lobes distinctly longer than high. Eli-
maea poaefolia-gvoup 38
- Tegmen not so narrow; its width in middle dis-
tinctly longer than length of pronotum; costal
area usually widened in about basal half (figs.
158, 160, 164). Pronotum with lateral lobes only
little longer than high. Elimaeas. str 44
38. Ô S 39
- 9 9 41
39. Cerci distinctly shorter than subgenital plate.
Subgenital plate suddenly narrowed behind basal
area. West Sumatra E. (E.) jacobsonii
- Cerci of subequal length with subgenital plate.
Subgenital plate more gradually narrowed 40
40. Java E. (E.) poaefolia
- Borneo; also in Malaysia? E. (E.) rosea
41. Subgenital plate with lateral margins converging
in circa apical half; apex roundly excised and with
apico-lateral angles acute. Java. ....£. (E.) poaefolia
- Subgenital plate with apex not roundly excised ...
42
42. Fastigium verticis not furrowed. Subgenital plate
short, apex obtuse, at each side provided with a
narrow, acute lobe. Borneo; also in Malaysia?
E. (E.) rosea
- Fastigium verticis furrowed. Subgenital plate
without a narrow acute lobe at each side. Thailand
43
43. Subgenital plate with apex transversely truncate
and slightly excised in middle; at each apico-later-
al angle with 2 short obtuse lobes (fig. 115). Go-
nangulum of ovipositor longer, pointing more or
less apicad (fig. 138). Northern Thailand
E. (E)sp. 1
- Subgenital plate with apex obtuse-triangular with
somewhat irregular margin, at each side with a
short conical projection (fig. 114). Gonangulum
of ovipositor shorter, pointing more or less ven-
trad (fig. 137). Western Thailand E. (E.)sp. 2
44. Male: Subgenital plate split in circa apical half or
more (figs. 55-56). Cerci rather strongly bulging
before apex (figs. 49-50). Stridulatory file with
circa 28-29 rather large and spaced teeth (figs.
27-28). Female: Cerci with apex rather broad,
obtuse (fig. 145). Eastern Central Thailand,
Hawaii Islands E. (E.) nautica
- Male: Subgenital plate split in less than apical
half, usually split in apical third only (figs. 59-
65); if split for almost apical half (figs. 57-58)
then stridulatory file with 34-42 teeth (figs. 24-
25). Cerci only slightly widened before apex (figs.
42-48). Female: Cerci with apex narrow, suba-
cute (figs. 146-150) 45
45. Male: Stridulatory file with less than 20 large and
widely spaced teeth in basal and central area and
a variable number of minute teeth in apical area
(figs. 18-23). Female: Gonangulum of ovipositor
with a large ventro-apical appendage (figs. 127-
128). Apex of subgenital plate only faintly excised
or apical lobes approached to each other (figs.
111-113) 46
- Male: Stridulatory file with more numerous and
less widely spaced teeth which are gradually de-
creasing in size from central area to apex (figs. 24-
26). Female: Gonangulum of ovipositor with a
small ventro-apical appendage (figs. 129-130).
Apex of subgenital plate with two widely spaced,
obtuse, apical lobes (fig. 110). [Female of E.
punctifera unknown] 47
46. Male: Stridulatory file with circa 32-35 (14-16
large) teeth. Female: Apex of subgenital plate
only faintly excised in middle (fig. 111). Cerci
less strongly curved (fig. 148). Central Thailand
to Java E. (E.) chloris
- Male: Stridulatory file with circa 9-17 (6-9 large)
teeth. Female: Apex of subgenital plate terminat-
ing into 2 subacute to subobtuse lobes which are
only narrowly separated from each other (figs.
112-113). Cerci more strongly curved (fig. 146).
Hongkong to North East India
E. (E.) subcarinata
47. Male: Stridulatory file with circa 53 teeth (fig.
26). Cerci with apical cone long and robust (fig.
44). Subgenital plate split in circa apical quarter
to apical third (figs. 61-62). Female unknown.
Bangladesh E. (E.) punctifera
- Male: Stridulatory file with circa 34-42 teeth
(figs. 24-25). Cerci with apical cone short and
73
Tijdschrift voor Entomologie, volume mi, 1998
slender (fig. 48). Subgenital plate split for some-
what less than apical half (figs. 57-58). Female:
Apex of subgenital plate with 2 short obtuse lobes
and broadly truncate in between (fig. 110). Thai-
land E. (E.) thaii
Elimaeini Yakobson, 1905: 324, 373
Elimaeini Yakobson, 1905: 324, 373; Bei-Bienko 1954: 115.
Elimaeae Brunner, 1878: 90.
Diagnosis. - Antennae elastic (less so in Ectadia).
Fastigium verticis forming a right angle with fastigi-
um frontis. Pronotum with a distinct humeral sinus.
Anterior coxa without spine but in some species with
a minute spinule. Anterior femur on ventral side fur-
rowed and provided with small spines. Anterior tibia
dorsally furrowed and provided with (small) dorso-
apical spurs. Tibial tympana covered by a conchate
fold with wide anterior slit on both sides or on inter-
nal (= anterior) side only. Tegmina fully developed,
hind wings surpassing tegmina in both sexes. Male
subgenital plate without styli. Ovipositor usually fal-
cate, rarely sabre-shaped, little longer than pronotum.
Ectadia Brunner, 1 878
Ectadia Brunner, 1878: 103. Type species: Ectadia pUosa
Brunner, 1878; by monotypy.
Diagnosis. - Elimaeini with modified tegminal ve-
nation and sexual wing dimorphism. Radius sector
fused with media; media with a zigzag course and
sending several oblique, subparallel branches to pos-
terior margin of tegmen. Tegmen of male with dorsal
area widened. In the female, both pairs of wings
slightly shortened but hind wings surpassing tegmina.
Tibial tympana conchate on both sides.
Distribution. - The genus is known from Kashmir
and from the mountainous region of eastern Burma,
northern Thailand and Yunnan (map 1).
Ectadia fulva Brunner
Ectadia fulva Brunner, 1893: 167. Holotype, a, Burma,
Carin Cheba, mcsn [not seen]. - Bei-Bienko 1962: 122.
Ectadia abbreviata Brunner, 1893: 167, fig. 58. Holotype,
9, Burma, Carin Ghecu, mcsn [not seen].
Material examined. — Thailand: 1 8 , Chiang Mai prov.,
Doi Suthep-Pui, 11 00- 1200m, 22.x. 1990; ló\ 19, same
data but 13.iv.1995; 23c?, 21 9, same data but ex ovo; 1 o\
1 9, Mae Hong Son prov., mountains between Samoeng -
Pa Pae, 30.iv.1988; 19, Nan prov., Doi Phukha, 1500m,
5.X.1991, alici.
Remarks. - The species is sufficiently described and
figured in Brunner (1893). The female of this species
was described under the name Ectadia abbreviata Brun-
ner, 1893 as already recognised by Bei-Bienko (1962).
The tegminal venation, especially the irregular
course of the media and the network of strong trans-
verse veinlets resembles the situation in Mirollia Stal,
1 873 but in a less extreme form. The rather stout and
breakable antennae and the sexual wing dimorphism
resemble the condition in the Actometopini. Howev-
er in contrast to the latter, the antennae are somewhat
more flexible and in the female, the hind wings still
surpass the tegmina although both pairs of wings are
slightly shortened. The male cerei are curved and
strongly compressed except for a short conical part at
base. They are similar to that of O. leeuwenii. E. fulva
shows a green brown colour dimorphism.
Development. - Oviposition was in parenchyma of
leaves of wheat, between the layers of absorbent paper
and in polystyrene. The eggs are of typical Phanero-
pterinae shape, compressed kidney-shaped. Egg de-
velopment required 37-94 (mean 52) days at 20-
23°C. There were 6 larval instars. The first larval
instar is pale reddish brown with 3 longitudinal white
stripes (fig. 168). Legs and antennae are almost colour-
less. Body and legs are dotted with black. Larval de-
velopment (from hatching to adult) required 52-68
days at 20-23°C. In captivity, larvae and adults readi-
ly fed the following European plants: leaves of Triti-
cum and Rumex as well as flowers of Rumex, Achillea,
Chrysanthemum, and Bellis, and less readily leaves of
Rubus and Taraxacum.
Stridulation (fig. 169). - The song of one male from
Doi Suthep (Chiang Mai province, Thailand) was
recorded at 20.5°C. Stridulation consists of a sequence
of about six echemes, the first echeme slightly quieter
than the following. Each echeme consists in the first
half of crescendoing pulses which are separated by
rather large pauses from each other. The second half of
an echeme is quieter, the repetition of the decrescen-
doing pulses becomes increasingly faster, such that in
the last quarter of an echeme single pulses can not be
recognised, even with a high time resolution. At the
end of an echeme the song intensity becomes loud
again. The echemes lasted between 1640 and 1890 ms,
the echeme interval varied between 4900 and 6050
ms. The main frequency of the song was about 6-14
kHz. The sound of an echeme can be circumscribed as
"trrrrr ziiip", with the second part of the sound quieter
than the first. Stridulation of Ectadia fulva with an ex-
treme acceleration of the stridulatory elements (sylla-
bles or pulses) within one echeme is rather curious.
Elir,
: Stal
Elimaea Stil, 1874: 27. Type species: Phaneroptera subcari-
nataStâï, 1861; by monotypy.
Diagnosis. - Elimaeini with posterior margin of
tegmen straight; radius, radius sector, media and cu-
74
Ingrisch: Review of Elimaeini
bitus running parallel and almost equally spaced from
each other; transverse veinlets dense, standing parallel
and almost vertical on principle veins (figs. 151-164).
Elimaea ( Orthelimaea) Karny
Orthelimaea Karny, 1926a: 23. Type species: Elimaea (Orthe-
limaea) leeuwen i i Karny, 1926a; by original designation.
Diagnosis. - Elimaeini with straight anterior femo-
ra. Tibial tympana covered by a conchate fold on both
sides. Greatest width of tegmen wider than length of
pronotum; radius sector branching about in middle
of tegmen. Male cerei either compressed or subcylin-
drical with an apical conus. Phallus membranous, but
dorsal lobes in some species provided with a longitu-
dinal rim covered with spinules. Ovipositor short
sabre-shaped with basal area substraight and apical
area with stout teeth or falcate with margins in apical
area finely serrulate; gonangulum with ventral margin
projecting, but sometimes not very distinct.
Remarks. - Orthelimaea is heterogeneous with re-
gard to the male cerei, the phallus and the female ovi-
positor. It is thus possible that this subgenus is a het-
erogeneous assemblage of species and must be further
subdivided. However, as most species referred to Or-
thelimaea are known from the Indian subcontinent, we
have to wait until those species are carefully revised.
Distribution. - The subgenus has its greatest species
diversity on the Indian subcontinent but spreads into
Central China and Thailand; a single species is known
from each, Java and Sulawesi (map 1).
Elimaea ( Orthelimaea) leeuwenii Karny
(figs. 2, 52, 78, 116, 119, map 3)
Elimaea {Orthelimaea) leeuwenii Karny, 1926a: 24, fig. 90.
Syn types, 26, Thailand, Bangkok, 5.xi.l920, Docters
van Leeuwen: 1 6 (without type label) in mzb [exam-
ined], 1 Ô (labelled holotype) in rmnh. - Ingrisch 1990a:
91, figs. 1-2.
Material examined. - Thailand: 29, Chiang Mai prov.,
Phrao district, Phrao - Ban Pradu, 26.-29.ix. 1985; 1 9 , Chi-
ang Mai prov., 8.x. 1991; 1?, Nakhon Ratchasima prov.,
Pak Chong, 1 6.x. 1990; 3d, Kanchanaburi prov., Thong
Pha Phum, 15.vi.1986; 19, Kanchanaburi prov., Erawan
falls, 17.-19.vi.1986; 2o\ 29, do. 2.vi.l988; 29, Prachuap
Khiri Khan prov., Hua Hin, 26.vii.1992, all ci.
Remarks. - The male of this species is already suf-
ficiently described in Karny (1926a), the female in
Ingrisch (1990a). The diagnostic characters are out-
lined in the key. The stridulatory file bears circa 31
teeth which are gradually decreasing in size towards
apex but remain rather large throughout.
The species is widespread in Central and Northern
Thailand and was so far collected from the Chiang
Mai province in the north to the Prachuap Khiri
Khan province in the south (map 3).
Development. - In contrast to all other Elimaea
species, E. leeuwenii has a sabre-shaped instead of a
sickle-shaped ovipositor. Oviposition of one female
in laboratory was in the soil; but the eggs were obvi-
ously not fertilised as no hatching occurred.
Elimaea (Orthelimaea) minor (Brunner) comb. n.
(figs. 3, 51,79, 151)
Elimaea minor Brunner, 1891: 48. Syntypes [not seen], 3
males: 26, Indonesia, Java, Tengger mountains, H.
Fruhstorfer (18.190; nhmw); \6, Java, Dr. Dohrn
(18.435; nhmw).
Elimaea (Rhaebelimaea) minor Karny 1926a: 20.
Material examined. - Indonesia: 1 6 , East Java, Tretes,
Gunung Arjuna, 1300m, mixed forest, 26.iii.1995; \6,
Central Java, Gunung Lawu, Tawangmangu - Sarangan,
1400m, mountain forest, 29.iii.1993, both ci.
Description. - Small species with parallel-sided
tegmen (fig. 151).
Male: Stridulatory file with circa 70 teeth which
are gradually decreasing in size towards apex (fig. 3).
Cerci regularly curved, slightly constricted in middle
and widened again before apex, apical cone subacute
(fig. 51). Subgenital plate slightly constricted behind
basal area, split into 2 lobes for more than apical half
(fig. 79); internal surface of resulting lobes densely
covered with minute spinules.
Female unknown.
Coloration: Green; posterior margin of pronotum
and dorsal field of tegmen (including stridulatory
area) brown. Tegmen with brown dots in cells be-
tween media and cubitus and a row of single larger
dots between radius (respectively radius sector) and
media. Variation: In the male from Gunung Lawu
which was collected as a freshly moulted specimen,
the dark pattern is largely absent.
Measurements of male (length in mm): Body 17;
pronotum 3.5; tegmen 21.0-23.0; tegmen width 4.0-
4.5; anterior femur 5.7-6.0; mesofemur 7.5-8.0; post-
femur 17.0-18.0.
Remarks. - In the original description (Brunner
1891) and later in Karny (1926a) it is said that this
species has the anterior femora curved in a phasmid-
like fashion. Accordingly, the species was combined
with Rhaebelimaea by Karny (1926a). This informa-
tion is based on an error. In the specimens at hand as
well as in the original type series (3d syntypes, "coll."
and "det. Brunner v. Wattenwyl" without type labels
in nhmw, information by Dr. A. Kaltenbach
19.xii.1996), the anterior tibiae are straight, without
the phasmid-like modification. The species must thus
be grouped with Orthelimaea.
75
Tijdschrift voor Entomologie, volume ui, 1998
Elimaea (Hemielimaea) Brunner stat. n.
Hemielimaea Brunner, 1878: 103. Type species: Hemieli-
maea chinensis Biunner, 1878; by monotypy.
Diagnosis. - Elimaeini with straight anterior femo-
ra. Tibial tympana covered by a conchate fold on in-
ternal, open on external side. Greatest width of
tegmen wider than length of pronotum; radius sector
branching about in middle of tegmen. Phallus with a
pair of conchate sclerites. Ovipositor falcate; gonan-
gulum with ventral margin projecting; dorsal margin
of ventral valves laterally projecting at base and
curved around projection of gonangulum.
Distribution. - The area of this subgenus stretches
from Eastern and Central China to Central Thailand
(map 1). There are probably more, superficially simi-
lar, Hemielimaea species in Thailand than the follow-
ing two. But the few additional specimens in dab (3 cT ,
39) and ci (1 9 ) do not allow to be certain about the
individual variation. The stridulato ry songs of individ-
uals from different localities should also be studied to
determine whether small morphological differences
are due to individual variation or are species specific.
Elimaea (Hemielimaea) procera (Ingrisch) comb. n.
(figs. 14-15,53,81,95, 118, map 3)
Hemielimaea procera Ingrisch, 1990a: 94, figs. 7, 14-16.
Holotype 6 : Thailand, Chanthaburi prov., Khao Soi
Dao, 3 l.v. 1986, smf [examined].
Material examined. - Thailand: 3$, Chanthaburi prov.,
Khao Soi Dao, 12.vi.1988, ci.
Description. - Fastigium verticis narrow, dorsal
margin sinuate, furrowed; step-like declined to fastigi-
um frontis. Pronotum with disc nearly flat, lateral
margins rounded; transverse sulcus broad V- or short
Y-shaped; paranota about as high as long; ventral and
posterior margins rounded together; humeral sinus
distinct. Tegmen sub-parallel-sided, in male about 6.7
times, in female 6.2-7.1 times longer than wide; ra-
dius sector branching in about middle. Anterior femur
with spines on both ventral margins. Meso- and post-
femur with spines on ventro-external margin only.
Knee lobes of all femora bispinose. Anterior tibia with
external and internal, dorsal and ventral apical spurs.
Male: Stridulatory file with a distinct step slightly
behind middle of length, with circa 100 teeth (figs. 14-
15). Tenth abdominal tergite with apex subtruncate.
Epiproct small, triangular. Cerci long, rather regularly
curved, hardly widened before apex, apical cone suba-
cute (fig. 53). Subgenital plate regularly curved from
base to apex, constricted before middle, split in circa
apical third, internal margins of resulting lobes dense-
ly covered with spinules (fig. 81). Phallus with a pair of
conchate sclerites with serrulate margin as in fig. 95.
Female: Tenth abdominal tergite narrow but entire.
Epiproct tongue-shaped. Cerci slightly curved, slight-
ly widening behind base, widest in circa middle, apex
subacute. Subgenital plate semicircular in general out-
line, apex subtruncate in middle; lateral areas faintly
grooved and with fine transverse furrows; central area
faintly and broadly furrowed (fig. 1 18); at each side of
subgenital plate intersegmental membrane with an
awl-shaped appendage. Ovipositor falcate, stout; dor-
sal margin serrate in circa apical half, ventral margin
in apical area; gonangulum slightly projecting ven-
trad, projection sticking in a curved fold of the dorsal
margin of the ventral ovipositor valves; dorsal margin
of the ventral ovipositor valves widened at base and
acute-angularly excised at end of widening.
Coloration: Green. Antenna black with spaced
light rings, but scapus and pedicellus with anterior
surface light, posterior (= dorsal) surface dark. Vertex
with a pair of longitudinal black bands. Pronotum
with complete or incomplete black lateral bands (dis-
tinct at anterior and posterior margins, often dis-
solved into dots or absent in middle). Tegmen with
dorsal area black with light veinlets, otherwise green
with dark dots in the cells between radius and cubitus
and between subcosta and anterior margin.
Measurements (length in mm): Body 3 21, 2 23-
26; pronotum S 4.5, 9 4.5-4.9; tegmen S 37.0, 9
39.0-40.5; tegmen width 6 5.5, 9 5.5-6.0; anterior
femur S 7.5, 9 8.5-9.0; mesofemur ó\ 9 11.5-13.0;
postfemur 6 26.0, 9 27.0-29.5; ovipositor 8.5-9.0.
Elimaea (Hemielimaea) cuculiata (Ingrisch) comb. n.
(figs. 16, 54, 80, 96, 1 17, map 3)
Hemielimaea cuculiata Ingrisch, 1990a: 96, figs. 8, 17-22.
Holotype 9: Thailand, Kanchanaburi prov., Erawan
falls, 17.-19.vi.1986, smf [examined].
Material examined. - Thailand: 2 S , 19, Kanchanaburi
prov., Erawan falls, 1. vi. 1988; 1 <3\ do. 9. iv. 1994 ex larva.
Description. - Fastigium verticis narrow, deeply
furrowed and dorsal margin sinuate; step-like de-
clined to fastigium frontis. Pronotum with disc near-
ly flat and lateral angles broadly rounded; anterior
margin concave, posterior margin convex; transverse
sulcus Y-shaped; paranota in male circa l.lx longer
than high, in female as high as long, ventral and pos-
terior margins rounded together; humeral sinus dis-
tinct. Tegmen in male 5.7-6.2 times, in female 5.8-
6.0 times longer than wide, anterior margin slightly
convex and tegmen thus gradually narrowing towards
apex; radius sector branching about in middle of
tegmen. Anterior femur with spines on both ventral
margins. Meso- and postfemur with spines on ventro-
external margin only. Knee lobes of all femora
bispinose. Anterior tibia with external and internal,
dorsal and ventral apical spurs.
76
Ingrisch: Review of Elimaeini
Male: Stridulatory file with a rather weak step in
circa middle of length, with about 90 - 109 teeth (fig.
16). Tenth abdominal tergite with a Y-shaped sulcus;
apical margin slightly sinuate. Epiproct long-triangu-
lar with apex broadly rounded. Cerci curved, slightly
widening behind base, widest in basal half, apex with
a strong, acute cone (fig. 54). Subgenital plate with
lateral margins sloping and approaching each other in
circa basal half, narrow and with subparallel lateral
margins in apical half; split in circa apical third, api-
cal lobes setose and on internal side of apical lobes
with strong spinules (fig. 80). Phallus with a pair of
conchate sclerites with serrate apical margin (fig. 96).
Female: Epiproct triangular, apex rounded. Cerci
slightly curved, widest in circa middle, apex obtuse.
Subgenital plate short-rectangular, apex rounded (fig.
117); at each side of subgenital plate intersegmental
membrane with an awl-shaped appendage. Ovipositor
falcate, stout; dorsal margin serrate in circa apical half,
ventral margin in apical area; gonangulum slightly
projecting ventrad, projection sticking in a curved fold
of the dorsal margin of the ventral ovipositor valves;
dorsal margin of the ventral ovipositor valves widened
at base and acute-angularly excised at end of widening.
Coloration: Green. Antenna black with spaced
light rings, but scapus and pedicellus with anterior
surface light, posterior (= dorsal) surface dark. Vertex
with a pair of longitudinal black bands. Pronotum
with incomplete black lateral bands (distinct at ante-
rior and posterior margins, dissolved into dots or ab-
sent in middle). Tegmen with dorsal area black with
light veinlets, otherwise green with indistinct dark
dots in the cells between radius and media and even-
tually between media and cubitus.
Measurements (length in mm): Body cT 20-22, 9
20-23; pronotum cT 4.2-4.5, 9 4.0-4.5; tegmen S
31.5-34.5, 9 36.0-38.0; tegmen width 6 5.5, 9 6.0-
6.5; anterior femur S 7.0-8.0, 9 7.5; mesofemur S
10.5-11.5, 9 11.0;postfemur S 22.0-24.5, 9 25.0;
ovipositor 8.0-8.5.
Stridulation (fig. 170). - The song of one male
from Erawan waterfall (Kanchanaburi province, Thai-
land) was recorded at 21.5°C. Stridulation consists of
an echeme with crescendoing syllables and, separated
by a short pause from the syllables, a loud final zip-
sound. The resulting sound can be circumscribed as
"trrrrrr zip", with the zip-sound louder than the first
part of the echeme. The echemes were often grouped
in loose sequences with 2-5 echemes which were re-
peated with an interval of 8-17 s. Between those se-
quences the male was quiet for several minutes.
The echemes lasted 1281-1443 ms. There were 16-
17 syllables per echeme. The syllables were repeated
with an interval of 55-84 ms, while the final zip-
sound followed 330-340 ms after the beginning of the
preceding syllable. Syllable duration varied between
21-73 ms. The main frequency of the song was about
13-24 kHz, that of the final zip-sound 9-24 kHz.
Stridulation of Hernielimaea cuculiata resembles
the songs of the Mediterranean Acrometopa species
(compare Heller 1988). This might be a hint for a re-
lationship between the Elimaeini and the Acrome-
topini as proposed by Brunner (1878), if the more
complex male stridulation of Hernielimaea compared
with Elimaea is the more primitive one. It is however
also possible that the similar time pattern in both gen-
era {Hernielimaea and Acrometopa) resulted from con-
vergent evolution.
Elimaea (Rhaebelimaea) Karny
Rhaebelimaea Karny, 1926a: 26. Type species: Pbaneroptera
parumpunctata Serville, 1839; here designated.
Diagnosis. — Elimaeini with anterior femora curved
as in phasmids. Tibial tympana covered by a conchate
fold on both sides. Greatest width of tegmen wider
than length of pronotum; radius sector usually branch-
ing about in middle of tegmen, rarely distinctly be-
fore middle. Phallus with a pair of conchate sclerites.
Ovipositor falcate, dorsal margin in apical half, ven-
tral margin only at apex finely serrulate; gonangulum
with ventro-apical angle projecting.
Remarks. - Several cases of misidentification of
species are summarised in Karny (1926a).
Distribution. - The species of this subgenus live in
a rather restricted area from Sumatra in the west to
the Philippines in the east and from Java in the south
to central Thailand in the north (map 2). A few
species from South India and Sri Lanka might also
belong here, but without re-examining the speci-
mens, it is not possible to be certain about their sub-
generic status. Rhaebelimaea species usually live in
forests and have very restricted ranges, many of them
have so far been recorded from a single locality.
Elimaea (Rhaebelimaea) parumpunctata (Serville)
(figs. 6, 32, 68, 87, 97, 121, 136, 152, 167)
Pbaneroptera parumpunctata Serville, 1839: 418. Holorype,
9 , Indonesia, Java, mnhn? [not seen].
Elimaea {Rhaebelimaea) parumpunctata Karny 1926a: 34,
fig. 95.
Material examined. - Indonesia: 1 $ , West Java, Gunung
Salak, above Ciapus, Sukamandri, 850-950m, mountain for-
est, 27. ii. 1995; 4<3\ same data but ex ovo; 1 $ , West Java,
Cibodas, Botanical Gardens, 1400m, 24.iii.1993 ex larva;
1 2 , same data but 1 .iv. 1 995, all ci.
Description. - Male: Stridulatory file with circa 104
teeth which are gradually decreasing in size towards
apex (fig. 6). Cerci shorter than subgenital plate, strong-
ly curved, widened before apex, apical cone slender,
subacute (fig. 32). Subgenital plate strongly narrowing
77
Tijdschrift voor Entomologie, volume i4i, 1998
from base to circa middle of length, apical area nar-
row; apex split into 2 slightly diverging lobes with
obliquely truncate apex (fig. 68). Phallus with a pair of
semi-ovoid, conchate sclerites (fig. 87).
Female: Subgenital plate transverse, subdivided in
middle, apex with slightly diverging cones at lateral
angles, sinuate in between (fig. 97). Ventral oviposi-
tor valves with a tongue-shaped appendage at base be-
tween gonangulum and subgenital plate (figs. 121,
136); gonangulum with a weak, broadly rounded
ventral projection (fig. 136).
Coloration: Green. Pronotum with incomplete black
lateral bands and with black dots. Tegmen in male with
dorsal field black with orange veinlets, in female
green with black dots; aggregations of black dots in
cells between media and cubitus and rows of single
larger dots between radius (respectively radius sector)
and media. Variation: In the female the dark pattern
is hardly expressed; instead, in the live specimen,
pronotum with white lateral bands and tegmen with
white posterior margin (fig. 152).
Measurements (length in mm): Body 3 21-22, 9 20-
21; pronotum 3 3.8-4.2, 9 3.5-3.8; tegmen 3 31.0-
32.0, 9 31.0-33.0; tegmen width 3 5.0, 9 5.0-5.5; an-
terior femur 3 7.0, 9 7.5-8.0; mesofemur 3 9.5-10.0,
9 9.0-10.0; postfemur 3 20.5-22.0, 9 21.0-22.0.
Development. - Oviposition was in parenchyma of
leaves. The eggs are compressed kidney-shaped. Egg
development required 41-43 days at 25-28°C. There
were 6 larval instars. The first instar larvae is green
with 3 longitudinal rows of white dots and the ab-
dominal apex white (fig. 167). The legs are olivaceous
to yellowish brown and densely covered with black
dots. The antennae are blackish brown with white an-
nulation. Larval development (from hatching to adult)
required about 63-75 days (temperature not controlled,
but always above 20°C). In captivity, the species fed on
a variety of plants including Mentha, Rumex, wheat,
seedlings of sunflower, as well as on dry rolled oats.
Stridulation (fig. 171). - Stridulation of two males
bred from the egg from Gunung Salak (West Java, In-
donesia) was recorded at 24.5° and 27°C. Two differ-
ent songs were produced by the same male, a short
song consisting of a single syllable and a long song
consisting of a sequence of 4-8 syllables. The time pat-
tern of the syllables was the same in both song types.
The syllables lasted 29-73 ms in the long songs at
27°C and 54-61 ms in the short songs at 24.5°C. The
long songs lasted 1210-2500 ms, depending on the
number of syllables, with a syllable repetition rate of
282-312 ms. The mean frequency of both song types
was 12-24 kHz. The sound of a syllable can be cir-
cumscribed as "zip", the long songs are a repetition of
this sound.
Elimaea {Rhaebelimaea) curvicercata Brunner stat. rev.
(figs. 11,33,66,88, 155)
Elimaea curvicercata Brunner, 1891: 50. Syntypes, a, 9,
Java orientalis, nhmw [not seen] .
Elimaea {Rhaebelimaea) parumpunctata partim Karny
1926a: 34.
Material examined. - Indonesia: 1 â , East Java, Tretes,
Gunung Arj una, 1000m, mixed forest, 26.iii.1995, ci.
Description. - Male: Stridulatory file with circa 79
teeth, gradually decreasing in size towards apex (fig.
11). Cerci strongly and angularly curved, distinctly
constricted and then widening again before apex, api-
cal cone long, apex subacute (fig. 33). Subgenital plate
narrowing from base to circa middle of length, apical
area narrow, apex split into 2 lobes with bulging inter-
nal margin, apex of lobes obliquely truncate (fig. 66).
Phallus with a pair of very slender, conchate sclerites
(fig. 88).
Female: According to the description in Brunner
(1891), the subgenital plate is similar to that of R.
parumpunctata, but the medial lobe emarginate in
middle.
Coloration (male): Green. Pronotum with black
lateral bands which are obsolete in metazona and with
black dots. Tegmen with dorsal field black with or-
ange veinlets, stridulatory vein green; aggregations of
black dots in cells between media and cubitus and
rows of single larger dots between radius (respectively
radius sector) and media. Mesofemur with a ventral
black spot at base and a row of black dots on external
surface. In the live specimen, central part of vertex,
disc of pronotum and anterior part of stridulatory
area of tegmen before stridulatory vein white.
Measurements of male (length in mm): Body 19;
pronotum 3.8; tegmen 29.0; tegmen width 6.0; ante-
rior femur 6.0; mesofemur 9.0; postfemur 20.0.
Remarks. - The species was previously regarded to
be a synonym of E parumpunctata (Serville, 1839)
(Karny 1926a). It shows however distinct differences
in the stridulatory file as well as in external and inter-
nal male genitalia. It is regarded here as a distinct
species. The differences between both taxa are out-
lined in the key.
Elimaea (Rhaebelimaea) kraussi Karny
(figs. 10, 17, 34, 67, 92, 105)
Elimaea (Rhaebelimaea) kraussiY^my, 1926a: 31, fig. 92. Syn-
types, 4o\ 1 9, West Java, Cibodas, 1400m, xi./xii.l921,
H.H. Karny, 3c? in mzb [examined], 1 S , 1 9 in rmnh.
Material examined. - Indonesia: IS , West Java, Gunung
Pangrango, Botanical Garden to Cibeureum waterfall,
1400-1700m, primary mountain forest, 25.iii.1993, ci.
Description. - Male: Stridulatory file with circa
78
Ingrisch: Review of Elimaeini
102 teeth which are distinctly spaced in circa basal
half (circa 35 teeth), very narrow and dense in apical
half; at apex there are a few small and widely spaced
teeth (fig. 10). The dorsal field of the right tegmen
(covered by the left tegmen when at rest) carries a se-
ries of rather regularly spaced transverse veinlets
which are raised and forming transverse ridges (fig.
17). Cerci short, substraight at base and then curved
in a more than 90 "-angle before the very long and
slender apical cone (fig. 34). Subgenital plate with
apical area narrow, apex shortly divided into 2 lobes
with faintly bulging internal margins, apex of lobes
obliquely truncate (fig. 67). Phallus with a pair of cir-
ca semi-ovoid conchate sclerites (fig. 92).
Remarks. - The species was already extensively de-
scribed by Karny (1926a). Some additional notes on
stridulatory file and male genitalia are added.
Elimaea (Rhaebelimaea) mentaweii sp. n.
(figs. 7, 40, 77, 84-86, 100, 123, 135, 144, 159)
Holotype 6: Indonesia, Mentawei exp., H.H.
Karny, R.M. 17.x. 1924, mzb. - Paratypes: Indonesia,
Mentawei exp., H.H. Karny, all mzb: IcT, no. 95,
Siberut, 23.ix.1924; 1 9, no. 112, Siberut, 25.ix.1924;
la, no. 163, Sipora, 9.x. 1924; 2â, no. 170, Sipora,
10.x. 1924; 12, no. 173, Sipora, 11.x. 1924; 2 9, no.
174, Sipora, 1 2.x. 1 924; 1 9, no. 183, Sipora, 15.X.1924;
l<?,no. 187, 17.x.l924;2 2,no. 188, 17.x. 1924; 19,
no. 227, Sipora, 3 1.x. 1924; 26, R.M. 17.x. 1924; 1 6,
19, RM. 18.x. 1924; 16, RM. 2 1.x. 1924; 19 (allo-
type), RM. 22.X.1924; là, Mentawei, Sipora, Sere-
ina, v.-vi.l894, Modigliani, mcsn; 36 , 1 9 , Mentawei,
Si Oban, iv.-viii.1894, Modigliani, mcsn.
Description. - Fastigium verticis with dorsal surface
roundly declined; dorsal surface shallowly furrowed.
Pronotum with disc slightly rounded to nearly flat,
but in posterior area slightly concave, lateral margins
rounded into paranota; anterior margin concave in
middle, posterior margin rounded; transverse sulcus
circa triangular, in or slightly behind middle of prono-
tum length; paranota about as long as high or faintly
longer, anterior angle angularly rounded, posterior an-
gle broadly rounded, humeral sinus weak but distinct.
Tegmen surpassing hind knees; with subparallel trans-
verse veinlets in circa basal half of tegmen; radius sec-
tor branching somewhat before or behind middle of
tegmen or even distinctly behind, this character some-
what variable even on left and right tegmen of the
same specimen. Anterior femur with dorsal margins
angular. Anterior femur with spines on ventro-inter-
nal margin or on both ventral margins (varying be-
tween individuals), mesofemur with spines on ventro-
external margin, postfemur without spines or with a
few spines on ventro-external margin. Knee lobes of all
femora bispinose or individual lobes unispinose.
Male: Stridulatory file with circa 82 teeth which
are distinctly spaced in basal half, dense but regularly
spaced in apical half (fig. 7). Tenth abdominal tergite
with apical margin subtruncate. Epiproct long tongue-
shaped, slightly curved from base to apex, shallowly
depressed in basal area (fig. 84a). Cerci long, curved
ventrad near base and mediad near apex, at the turn-
ing point between ventral and medial curvature cer-
cus slightly narrowed; apex forming a curved, tri-
edged cone (fig. 40). Subgenital plate elongate, basal
area with sloping lateral margins, central area pro-
longed behind and narrow, apex split to a variable de-
gree with apices of lobes obtuse (fig. 77). Phallus with
a pair of mussel-shaped, large, hyaline sclerites, with
concave internal and convex external surface; external
surface in basal area covered by membrane; dorso-api-
cal margin finely serrulate (figs. 85-86).
Female: Tenth abdominal tergite with apical mar-
gin subsinuate. Epiproct long, triangular with apex
rounded; dorsal surface slightly curved from base to
apex. Cerci curved, widest near middle of length; apex
pointed (fig. 144). Subgenital plate transverse, divided
in midline but both halves connected by strong mem-
brane, apico-lateral angles acute-angularly projecting
behind (fig. 100). Gonangulum of ovipositor with a
short, obtuse, ventral projection that inserts into a
fold of the ventral ovipositor valves (fig. 123, 1 35).
Coloration: Green when alive. Pronotum: disc
margined with dark brown lateral bands and dark
spots on both sides of the bands (more distinct incT
than in 9). Tegmen: area behind media with dark
cells and pale veinlets; area between radius and media
usually with one or behind branching of radius sector
with 2 rows of dark spots; towards apex with or with-
out small dark dots; dorsal field brown in male only.
Measurements (length in mm): Body 6 18-23, 9
17-23; pronotum cT 4.0-5.0, 9 4.5-4.7; tegmen cT
35.0-40.0, 9 37.0-41.0; tegmen width 6 6.0, 9 6.0-
6.5; postfemur 6 24.0-29.0, 9 27.0-29.0; ovipositor
8.0-8.5.
Etymology. - The name of the new species is cho-
sen from the type locality, the Mentawei Islands west
of Sumatra.
Elimaea (Rhaebelimaea) maninjauensis sp. n.
(figs. 8, 39, 76, 94, 99, 122, 143, 153)
Holotype 6 : Indonesia, West Sumatra, Maninjau,
500-700m, I6.iii.1995 ex larva, S. Ingrisch, mzb. -
Allotype: 1 9 , Indonesia, West Sumatra, Maninjau,
I4.iii.1993, ci.
Description. - Fastigium verticis narrow, sulcate,
apex obtuse and step-like declined to fastigium fron-
ds. Pronotum with disc nearly flat, especially in pos-
79
Tijdschrift voor Entomologie, volume ui, 1998
terior area, lateral angles rounded, apical area shoul-
dered; anterior margin faintly concave in middle, pos-
terior margin rounded; transverse sulcus Y-shaped;
paranota almost as high as long; ventral margin round-
ed, humeral sinus present. Tegmen surpassing hind
knees; anterior margin convex, posterior margin sub-
straight; radius sector branching slightly before or in
middle of tegmen. Anterior femur with dorsal mar-
gins angular. Anterior femur with spines on ventro-
internal margin, mesofemur with spines on ventro-
external margin, postfemur without spines or with
one spine on ventro-external margin. Knee lobes of
anterior femur bispinose on external, unispinose on
internal side, of mesofemur unispinose on external,
bispinose on internal side, of postfemur bispinose on
both sides. Anterior tibia with dorsal and ventral, ex-
ternal and internal, apical spurs.
Male: Stridulatory file curved behind base, other-
wise substraight with about 1 26 teeth which become
regularly narrower from base to apex (fig. 8). Tenth
abdominal tergite distorted in specimen at hand. Epi-
proct long tongue-shaped, apex rounded, with a broad
medial furrow in basal area. Cerci strongly curved near
base, afterwards moderately curved; apex terminating
into a stout, triangular, acute tooth (fig. 39). Subgen-
ital plate narrow even at base, basal area with sloping
lateral margins, central area prolonged behind into a
long, narrow, medial process with subparallel margins
behind middle and with a weak and broad medial fur-
row, apex shortly divided into two obtuse lobes (fig.
76). Phallus with a pair of mussel-shaped, large, hya-
line sclerites, with serrulate margin (fig. 94).
Female: Tenth abdominal tergite with apical mar-
gin subtruncate. Epiproct triangular, apex rounded;
dorsal surface slightly curved from base to apex. Cer-
ci very thin-cylindrical, moderately curved, apex spin-
ose (fig. 143). Subgenital plate transverse, in midline
with a membranous fold, apical margin subtruncate
or slightly concave and on each apical angle pro-
longed into a long spinose projection (fig. 99). Go-
nangulum of ovipositor with a short obtuse ventro-
apical projection pointing ventrad.
Coloration: Green. Pronotum with narrow, dark,
lateral bands accompanied by dark dots. Tegmen with
black dots and with two rows of spaced larger dots in
radial areas. In male, tegmen with dorsal area brown.
Anterior femur with black spots around spines on ven-
tral edges.
Measurements (length in mm): Body S 17, 9 27;
pronotum S 4.5, 9 4.5; tegmen S 35.0 (broken), 9
39.0; tegmen width 6 6.0, 9 7.0; postfemur S 23.0,
9 25.0; ovipositor 7.5.
Etymology. — The name of the new species is cho-
sen from the type locality, Lake Maninjau in West
Sumatra.
Elimaea (Rhaebelimaea) modiglianii sp. n.
(figs. 12-13, 31, 72, 89, 102, 132, 140)
Holotype 6: Indonesia, Sumatra, Si - Rambe,
xii.1890 - iii.1891, E. Modigliani, mcsn. -Paratypes:
Indonesia: 2<S, Sumatra, Si - Rambe, xii.1890 -
iii.1891, E. Modigliani, mcsn; 1 9 , Balighe, x.1890 -
iii.1891, E. Modigliani, mcsn.
Description. - Fastigium verticis narrow, sulcate,
apex obtuse, step-like declined to fastigium frontis.
Pronotum with disc narrow, subparallel-sided, surface
nearly flat, in anterior area rounded; lateral angles
rounded into paranota, apical area shouldered; trans-
verse sulcus V- or Y-shaped; paranota slightly longer
than high in male or about as long as high in female;
ventral margin rounded, humeral sinus present.
Tegmen surpassing hind knees; anterior margin con-
vex, posterior margin substraight; radius sector branch-
ing slightly behind middle of tegmen. Anterior femur
with dorsal margins angular, with spines on both ven-
tral margins, those on internal side sitting on dark
brown spots; meso- and postfemora with spines on
ventro-external margin only (in the female with a sin-
gle spine also on ventro-internal margin of postfemur).
Knee lobes of all femora bispinose, but the ventral
spine often smaller and occasionally absent (especially
on external side of postfemur). Anterior tibia with dor-
sal and ventral, external and internal, apical spurs; dor-
so-internal spur smaller and occasionally absent.
Male: Stridulatory vein widened and greatly bulging
on dorsal side. Stridulatory file step-like declined be-
fore middle (after about 22 teeth), teeth large and
wide before the step; the step includes about 7 teeth;
behind the step with about 62 - 67 teeth which are be-
coming smaller and denser towards apex (figs. 12-13).
Tenth abdominal tergite transverse with apex subtrun-
cate. Epiproct long tongue-shaped, apex triangularly-
rounded, with a broad and shallow furrow in basal
area. Cerci rather regularly curved; apical area narrow-
spatulate, apex acute (fig. 31). Subgenital plate strong-
ly curved dorsad behind middle of length; with a weak
medial carina; apical area roundly excised, with lobes
narrowly spaced and subparallel, apex of lobes trans-
versely truncate (fig. 72); divided apical area provided
with long hair on dorsal, internal and lateral surfaces.
Phallus with two elongate conchate sclerites (fig. 89).
Female: Tenth abdominal tergite with apical mar-
gin subtruncate. Epiproct tongue-shaped, apex broad-
ly subtruncate. Cerci rather stout, regularly curved
and narrowed to apex, apex subobtuse (fig. 140). Sub-
genital plate short, much wider than long, with a
weak median keel; apico-lateral angles acute-triangu-
larly projecting; central part of apical margin subsin-
uate (fig. 102). Gonangulum of ovipositor with a
short, obtuse, ventral projection (fig. 132).
80
Ingrisch: Review of Elimaeini
Coloration: Probably green when alive; specimens
at hand pale brown (discoloured during previous stor-
age in alcohol). Pronotum with narrow black bands
bordering disc and some dark dots on paranota below
those bands. Tegmen with dorsal field (including
stridulatory area) dark brown, but veins and veinlets
pale; stridulatory vein black; with aggregations of
dark dots in cells of cubital area and single dark dots
in some cells of medial area.
Measurements (length in mm): Body â 14-16, 9
19; pronotum cT 4.0, 9 4.0; tegmen cT 33-36, 9 32;
postfemur cT 23.0-26.0, 9 22.0; ovipositor 5.8.
Etymology. - The name of this species is given af-
ter the collector, Dr. E. Modigliani.
Elimaea (Rhaebelimaea) transversa Ingrisch
(figs. 98, 133, 141)
Rhabelimaea [sic] transversa Ingrisch, 1990a: 91, fig. 3-6.
Holorype 9: Thailand, Chanthaburi prov., Khao Soi
Dao, 15.x. 1985, smf [examined].
Material examined. - 1 9, Thailand, Chanthaburi prov.,
Khao Soi Dao, 12.vi.1988 (ci); 19, Chanthaburi prov.,
Pong Nam Lorn, 30.iv.1959, Pai San (dab, Lot 2506).
Measurements of female (length in mm): Body 22-
23; pronotum 4.0-4.8; tegmen 34.0-35.0; tegmen
width 7.0-7.5; postfemur 23.5-24.0; ovipositor 7.0-7.5.
Remarks. - The species is already sufficiently de-
scribed in Ingrisch (1990a). It was previously only
known from the holorype. New material available in-
cludes two females; thus the male is still unknown.
The diagnostic characters are outlined in the key.
Elimaea (Rhaebelimaea) pseudochloris sp. n.
(figs. 9, 36, 73, 93, 161)
Holotype 6: Thailand, Trang prov., Khao Chong,
23.x. 1991, at light, S. Ingrisch, dab.
Description. - Fastigium verticis narrow, sulcate,
apex obtuse, step-like declined to fastigium frontis.
Pronotum with disc nearly flat and with a distinct
medial carina which is subobsolete in posterior area;
paranota longer than high, ventral margin subsinuate
and angles rounded, humeral sinus distinct. Anterior
femur with spines on both ventral margins, meso-
and postfemur with spines on ventro-externaJ mar-
gin. Knee lobes of all legs uni- or bispinose (irregular).
Anterior tibia with dorsal and ventral, external and
internal, apical spurs. Tegmen surpassing hind knees,
anterior margin distinctly convex, posterior margin
faintly concave, substraight; radius sector branching
in basal half, distinctly before middle of tegmen.
Male: Stridulatory file with circa 32 teeth which are
rather large and almost equally sized and equally spaced
throughout (fig. 9). Tenth abdominal tergite with api-
cal matgin subtruncate (faintly concave). Epiproct long
tongue-shaped. Cerci narrow-cylindrical, strongly
curved but substraight at base; apex terminating in a
long twisted cone with acute apex (fig. 36). Subgenital
plate with a short bowl-shaped basal part and a long,
curved, apical part which is parallel-sided and split
from apex for slightly less than half of its length (fig.
73); resulting lobes armed with numerous spinules on
medial surfaces. Phallus with large semi-cylindrical
projections which are terminating in mussel-shaped
sclerites with apical margin finely serrulate (fig. 93).
Female unknown.
Coloration: Green; pronotum, legs and body with
black dots. Pronotum with a yellow medio-longitudi-
nal band; abdominal tergites red in middle.
Measurements of male (length in mm): Body 20;
pronotum 4.0; tegmen 29.5; tegmen width 5.5; post-
femur 21.5.
Etymology. - The name of this species is given for
its superficial similarity with E. chloris.
Elimaea (Rhaebelimaea) apicata sp. n.
(figs. 4, 37, 74, 91, 154)
Holorype 6, Thailand, Surat Thani province,
Khao Sok, 150m, 23.L1997, S. Ingrisch, dab.
Description. — Fastigium verticis narrow, dorsally
furrowed; in lateral view slightly sinuate; step-like de-
clined to fastigium frontis. Pronotum with disc flat-
tened, with indication of lateral carinae and rounded-
angularly bent into paranota; lateral margins subparallel,
only at apex slightly diverging; anterior margin slightly
concave, posterior margin rounded but somewhat
constricted in middle; transverse sulcus short Y-
shaped, with indication of a short medial carinula be-
fore apex; paranota slightly longer than high (l.lx);
ventro-posterior angle broadly rounded, humeral si-
nus distinct. Tegmen with anterior area faintly
widened in basal half, with almost subparallel mar-
gins, apex rounded; radius sector branching in about
middle (15 mm from base). Anterior femur distinctly
compressed with spines on both ventral margins.
Meso- and postfemur with spines on ventro-external
margin. Genicular lobes of all legs bispinose. Tibial
tympana conchate but leaving a father large propor-
tion of the tympana uncovered. Anterior tibia with
dorsal and ventral, external and internal apical spurs.
Male: Stridulatory file with circa 90 rather dense
teeth which gradually become narrower from base to
apex (fig. 4). Tenth abdominal tergite largely pro-
longed behind and divided into two rounded lobes in
apical third. Epiproct tongue-shaped, covered by the
projection of tenth tergite. Cerci long, in basal area
substraight, afterwards strongly curved, apical area
81
Tijdschrift voor Entomologie, volume 141, 1998
compressed and terminating in a long-triangular,
acute cone (fig. 37). Subgenital plate with quickly ap-
proaching lateral margins in basal area, central area
attenuate and prolonged, gently curved and sub-par-
allel-sided, only at apex very faintly excised; basal part
with a medial carina that runs to about half of the at-
tenuate central projection (fig. 74). Phallus with a pair
of elongate, sclerotised, conchate projections with set-
rate dorsal margin (fig. 91).
Female unknown.
Coloration: Green. Body and legs with black and
red dots. Antenna brown on dorsal, light on ventral
side, in posterior area blackish-brown with largely
spaced white rings. Pronotum with lateral carinulae
whitish, bordered on both sides by a band of black
dots. Tegmen green, dorsal field black but with stri-
dulatory vein, a large spot at end of stridulatory area
and veinlets light brown; lateral area with 2 rows of
large black dots (between subcosta and anterior mar-
gin and between radius and media) and with aggrega-
tions of small black dots in cells between radius and
anal margin.
Measurements of male (length in mm): Body 22;
pronotum 3.8; tegmen 31.5; tegmen width 4.8; ante-
rior femur 7.5; mesofemur 1 1.5; postfemur 22.5-
Etymology. - The name of this species refers to the
prolongation of the last abdominal tergite.
Elimaea (Rhaebelimaea) sinuata sp. n.
(figs. 5, 30, 71,90, 162)
Holotype 6: Indonesia, Mentawei exp., H.H.
Karny, Sipora no. 195, 2 1.x. 1924, mzb.
Description. - Fastigium verticis narrow, furrowed
above; step-like declined to fastigium frontis. Prono-
tum with disc nearly flat especially in posterior area,
angles rounded, anterior margin concave, posterior
margin rounded, transverse sulcus V-shaped; paranota
circa as long as high, humeral sinus present. Tegmen
surpassing hind knees, sub-parallel-sided but faintly
curved throughout, apex rounded; radius sector
branching slightly before middle; transverse veinlets
rather regular, especially in anterior area and between
radius and radius sector. Anterior coxa with a minute
spinule. Anterior femur with one spinule on both ven-
tral margins, meso- and postfemur with several spin-
ules on ventro-external margins. Knee lobes of all
femora bispinose. Tympana conchate on both sides,
but leaving the anterior part of the tympanum uncov-
ered. Anterior tibia with dorsal and ventral, external
and internal, apical spurs. Posttibia with one dorsal
and two ventral apical spurs on both sides.
Male: Stridulatory file sinuate, with about 10 large,
widely spaced teeth in circa basal half and narrow,
dense teeth in apical half; together about 52 teeth (fig.
5). Cerci long-cylindrical, rather moderately curved,
apex slightly bulbous and with a large, compressed,
curved, subacute cone (fig. 30). Subgenital plate bowl-
shaped, regularly curved throughout, split until base,
apex terminating in two narrow, parallel, subacute,
stylate projections (fig. 71). Phallus with a pair of
wedge-shaped sclerites with slightly rounded apical
margin curved ventrad and serrulate; and ventral of
sclerites with an unpaired stylate projection which is
slightly curved and has angular margins, dorsal angles
armed with spines (fig. 90).
Female unknown.
Coloration: Yellowish brown, probably green when
alive. Pronotum with interrupted bands of spots and
dots along lateral angles. Tegmen with a row of dark
spots in radial area.
Measurements of male (length in mm): Body 19;
pronotum 4.8; tegmen 35.0; tegmen width 5.5; post-
femur 26.0.
Etymology. - The name of this species refers to the
sinuate course of the stridulatory file.
Elimaea (Rhaebelimaea) pentaspina sp. n.
(figs. 101, 124, 134, 142)
Holotype 9: Thailand, Chanthaburi prov., Khao
Soi Dao, 19.xii.1974, A.Lewvanich, dab (Lot 3697).
- Paratype: 1 9, Thailand, Chanthaburi prov., Khao
Soi Dao, 12.vi.1988, ci.
Description. - Fastigium verticis narrow, sulcate,
apex obtuse and step-like declined to fastigium frontis.
Pronotum with disc broadly rounded but posterior
area nearly flat and shouldered, lateral angles rounded;
anterior margin concave in middle, postetior margin
rounded; transverse sulcus Y-shaped, behind middle
of pronotum; paranota longet than high, ventral mar-
gin rounded, humeral sinus weak. Tegmen surpassing
hind knees; radius sector branching slightly before
middle of tegmen. Anterior femur phasmid-like curved
but curvature weak, almost straight. Anterior femur
with spines on both ventral margins, meso- and post-
femur with spines on ventro-external margin. Knee
lobes of all femora bispinose. Anterior tibia with dor-
sal and ventral, external and internal, apical spurs.
Male unknown.
Female: Tenth abdominal tergite with apical margin
subtruncate. Epiproct long tongue-shaped. Cerci long-
conical, curved, apex obtuse (fig. 142). Subgenital
plate with a compressed lateral projection at each side;
apical margin with a long spinose projection at each
angle and area in between short, obtuse-angularly pro-
jecting (fig. 101). Gonangulum of ovipositor with a
long, conical, ventro-apical projection; ventral oviposi-
tor valves with a triangular projection with obtuse apex
just below projection of gonangulum (figs. 124, 134).
82
Ingrisch: Review of Elimaeini
Coloration: Green; abdomen with red dots. Prono-
tum with black dots especially around lateral angles.
Tegmen with a longitudinal band of black dots in
medial area and some dots in other fields.
Measurements of female (length in mm): Body 21-
23; pronotum 4.0-4.8; tegmen 34.0-34.5; tegmen
width 6.0; postfemur 24.0-25.0; ovipositor 6.5-6.8.
Etymology. - The name of this species refers to the
subgenital plate with its 5 (2 lateral and 3 apical) pro-
jections.
Elimaea (Elimaea) Stai
Elimaea Stai, 1874: 27. Type species: Phaneroptera subcari-
nata Stil, 1861; by monotypy.
Diagnosis. - Elimaeini with anterior femora curved
as in phasmids. Tibial tympana covered by a conchate
fold on both sides. Greatest width of tegmen wider
than length of pronotum; radius sector branching dis-
tinctly before middle of tegmen. Phallus membra-
nous. Ovipositor falcate, dorsal margin serrulate in
circa apical half, ventral margin in apical area; gonan-
gulum with ventro-apical angle projecting.
Distribution. - The typical subgenus Elimaea has
its greatest species diversity in China and Indochina,
while from the Malay Peninsula to western Indonesia
there is only a single species (except for the poaefo-
/z'#-group). The known range spreads from the
Southern Maritime Territory of Russia in the north
to Java in the south, and from Taiwan in the east to
North-East India (West Bengal) in the west (map 1).
There are records of a few species with doubtful sub-
generic affinity from Bombay and Sri Lanka. Thus
the range of the subgenus might also cover the Indian
subcontinent. Species of the nominate subgenus are
often found in secondary vegetation, although they
can intrude into forests along roads and clearings.
Elimaea (Elimaea) subcarinata (Stal) stat. rev.
(figs. 20-23, 45-47, 59-60, 65, 112-113, 128, 146-
147, 158, 163, 164-165, map 3)
Phaneroptera subcarinata Stài, 1861: 319. Synrypes 1 d" , 1 $ :
China, Hongkong, nrs [examined].
Elimaea chloris (nee De Haan, 1842) Brunner 1878: 100
(partim); Karny 1926a: 36 (partim).
Elimaea appendiculata Brunner, 1878: 101. Holotype d:
Indochina, Thorey, nhmw (coli. Brunner no. 5503) [ex-
amined], syn. n.
Elimaea punetifera (nee Walker, 1869) Kirby 1906 (partim):
396; Uvarov 1927: 95 (partim).
Material examined. - Thailand: 1 d , 19, Chiang Mai
prov., Phrao district, Phrao - Ban Pradu, 26.-29.ix. 1985;
lo", 19, Lampang prov., Doi Khun Tan, 900m, 16
/17.ix.l993 at night; 26 d, 249, same data but ex ovo; ld,
Tak prov., Mae Salid, Monkrating, 700m, 18./19.V.1988;
ld, Nakhon Ratchasima prov., Khao Yai, 172.x. 1985, all
ei. - India: ld, West Bengal, Jalpaiguri, 18.xii.1986, R.S.
Barman; ld, 19, Mizoram, Teirei, Aizaul, 13.xi.1995,
M.S. Shishodia, all zsi.
Description. - Fastigium verticis narrow, apex suba-
cute, dorsal surface furrowed. Pronotum with disc flat-
tened, especially in posterior area, lateral angles round-
ed; transverse sulcus Y-shaped, slightly behind middle;
anterior margin concave, posterior margin rounded;
paranota longer than high (51:40 and 55:49 in syn types
of E. subcarinata, 56:48 in type of E. appendiculata);
ventral and posterior margins rounded; humeral sinus
distinct. Tegmen surpassing hind knees; radius sector
branching circa 8-10 mm from base. Anterior femur
with spines on both ventral margins (on external side
smaller). Meso- and postfemora with spines on ventro-
external margin. Genicular lobes of all legs bispinose.
Anterior tibia with external and internal, dorsal and
ventral apical spurs (dorso-internal spine lacking on one
fore leg of both syn types o£ E. subcarinata).
Male (syn type of E. subcarinata): Stridulatory file
(damaged) with 8 large to medium sized teeth and a
few indistinct minute teeth (fig. 20). Tenth abdominal
tergite with apex broadly rounded; central area setose
and with a small puncture in middle. Epiproct tongue-
shaped, apex obtuse. Cerci strongly curved; apical area
compressed, sinuate and external side convex, internal
concave, with narrowing margins and apex subacute
on dorso-cranial side (fig. 45). Subgenital plate moder-
ately curved dorsad (almost straight behind basal area);
split into two obtuse lobes in circa apical third (fig. 65).
Male (type of E. appendiculata): Stridulatory file
with 8 large and 3 minute teeth (fig. 21). Tenth ab-
dominal tergite entire (shrunk due to previous storage
in alcohol), central area setose. Epiproct tongue-
shaped with a weak medial carina. Cerci strongly
curved; apical area compressed, sinuate and external
side convex, internal concave, with narrowing margins
and apex acute on dorso-cranial side (fig. 46). Subgen-
ital plate damaged.
Male (specimens from Thailand): Stridulatory file
with 6-8 large and 2-11 minute teeth (fig. 22-23).
Tenth abdominal tergite with apex broadly rounded;
central area setose and with a small puncture in mid-
dle. Epiproct tongue-shaped, apex obtuse. Cerci
strongly curved; apical area compressed and twisted,
with narrowing margins; apex varying from acute (on
dorso-cranial end) to subobtuse (fig. 47). Subgenital
plate moderately curved dorsad; split into two obtuse
lobes in circa apical third (figs. 59-60).
Female: Tenth abdominal tergite with apex subtrun-
cate. Epiproct triangular, apex obtuse. Subgenital plate
acute-triangular, apex angularly excised and with 2
short triangular lobes (figs. 112-113). Cerci regularly
and rather strongly curved; apex subacute to subobtuse
(fig. 146, 147). Gonangulum of ovipositor with a large,
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Tijdschrift voor Entomologie, volume i4i, 1998
bulbous, ventro-apical appendage which in situ lies in a
cleft between dorsal and ventral valves (fig. 128).
Coloration: Syntypes of E. subcarinata: Discolored,
but probably green when alive. Antenna with scapus
and pedicellus concolorous, flagellum blackish brown
on dorso-lateral side (with antenna directed posteri-
orly), ventro-internal side light brown. Disc of prono-
tum in male with a brown medial band which is split
by a white line, in female green with the white line
only. Tegmen in cells between radius and anal margin
with aggregations of black dots, less distinct between
radius and media.
Type of £ appendi culata: Discolored, but probably
green when alive. Antenna with scapus and pedicellus
concolorous, flagellum dark brown especially on dor-
so-lateral side (with antenna directed posteriorly).
Tegmen in cells between media and anal margin, in
apical half also in cells between radius and media with
aggregations of black dots.
Variation of specimens from Thailand:
(1) Green. Compound eyes brown. Scapus and
pedicellus reddish brown, flagellum black in basal
area, with or without annulation, otherwise with
spaced light rings. Vertex and disc of pronotum with
a brown medial band. Tegmen green, anterior margin
orange or red, anal margin darkened (cells black,
veinlets green). Abdominal tergites red in middle; an-
terior and medial legs orange brown.
(2) as before but general colour yellowish brown
(this colour variant appeared while breeding the
species in laboratory). Antenna as described under (1)
or light brown with spaced dark rings.
(3) as (1) but dorsal area of tegmen distinctly dark-
ened.
(4) as (3) but anterior area of tegmen with a red
pattern; hind legs brownish.
(5) almost uniformly green (or yellowish brown);
flagellum of antenna only slightly infumate at base es-
pecially on dorsal side (with antenna directed posteri-
orly); vertex and pronotum without dark medial band;
tegmen green with the dark dots little conspicuous.
Measurements of syntypes of E. subcarinata (length
in mm): Body 6 16, 9 22; pronotum 6 4.2, 9 4.9;
tegmen S 29.0, 9 32.0; tegmen width S 4.7, 9 5.8;
anterior femur S 7.5, 9 9.0; mesofemur 0* 10.0, 9
10.0; postfemur S 22.0, 9 25.0; ovipositor 6.8.
Measurements of male type of E. appendiculata
(length in mm): Body 21; pronotum 4.5; tegmen
37.0; tegmen width 6.0; anterior femur 9.5; mesofe-
mur 12.0; postfemur 26.0.
Measurements of specimens from Thailand (length
in mm). - Body 6 21-28, 9 22-27; pronotum â
4.0-4.5, 9 4.2-4.8; tegmenc? 35.0-40.0, 9 35.0-
38.0; tegmen width S 5.5-6.0, 9 5.5-6.0; anterior
femur S 9.0-10.0, 9 9.5-10.0; mesofemur S 11.0-
12.5, 9 11.0-13.0; postfemur S 23.0-27.5, 9 23.5-
27.5; ovipositor 6.5-7.0.
Remarks. — Males of E. subcarinata are easily recog-
nisable by the low number of large and widely spaced
stridulatory teeth on the underside of the left tegmen.
The additional minute teeth at the apex which vary in
number and which are sometimes absent are certain-
ly not used for stridulation and as such do not under-
lie a selective pressure. The differences of the stridula-
tory files show with certainty that E. subcarinata is
not a synonym of E. chloris as supposed by Brunner
(1878), and E. appendiculata not a synonym of E.
punctifera as supposed by Kirby (1906).
The male syntype of E. subcarinata is rather small
compared with the series of specimens from Thailand
and with the type of E. appendiculata. Small differ-
ences in the male cerei might be attributed to this fact.
The stridulatory file does not show any significant dif-
ferences between the specimens of different origin.
Thus there is no doubt that they are conspecific, and
E. appendiculata is a synonym of E. subcarinata.
Development. - Oviposition was in parenchyma of
leaves {Rubus), but also between the layers of ab-
sorbent paper and in polystyrene. The eggs are com-
pressed kidney-shaped as typical for Phaneropterinae.
Egg development required 28-42 (mean 35) days at
25 °C and 39-105 (mean 64) days at 20-23 "C. There
are 6 larval instars. The first instar larvae are green
with 3 white longitudinal stripes which are very con-
spicuous (fig. 165). The femora are olivaceous at
base, otherwise legs and antennae are yellowish
brown. Larval development (from hatching to adult)
required 45-49 days at 25°C and 71-86 days at 20-
23°C. The spermatophore is very large (fig. 164). In
captivity, the species fed readily on European plants
as Rumex, Plantago, and seedlings of sunflower, and
less readily on wheat, Rubus, and Cirsium.
Stridulation (fig. 173). - The songs of two males
bred from the egg from Doi Khun Tan (Lampang
province, Thailand) was recorded at 24°C and 26°C.
Stridulation consists of loose groups of 1-3 syllables.
The groups are repeated at irregular intervals. Syllable
duration varied between 18-35 ms at 24°C or 15-26
ms at 26°C. It was thus similar to the syllable dura-
tion in E. chloris. However the syllables consisted of
only 4-6 pulses which were longer than in E. chloris
and not well separated from each other. Stridulation
was loudest at about 12-18 kHz. The sound of a syl-
lable can be circumscribed as "zip".
Elimaea (Elimaea) chloris (De Haan)
(figs. 18-19,42-43,63-64,83, 111, 120, 127, 148,
157, map 3)
Locusta (Phaneroptera) chloris De Haan, 1842: 192. Syn-
types (2cî, 1 $): Java, rmnh [examined]. - Elimaea chlo-
ris Brunner 1878: 100 (partim); Karny 1926a: 36, fig. 96
(partim); Karny 1926b: 265, fig. 185.
84
Ingrisch: Review of Elimaeini
Material examined. - Indonesia: 1 â , West Java, Palabuan
Ram, ii.1921, H.H. Karny, smf; 1 9, West Java, Bogor, Ke-
bun Raya, 15.ii.1995; North Sumatra, Pematang Siantar,
400m, 4.iii.l993. - Malaysia: 1 9, Pahang, Taman Negara,
Kuala Tahan, 19.-22.vii. 1984; 1 9, Pahang, Rantau Abang,
23.-25.vii. 1984. -Thailand: Id, Bangkok, Lard Phrao, 5.-
8.VÌ.1988; lo\ Surat Thani prov., Khao Sok, 150m,
24. i. 1997; 1 9 , Surat Thani prov., Koh Samui, Lamai beach,
10.X.1985; 29, do., Na Muang Falls, 25.ix.1989; 19, Pra-
chuap Khiri Khan prov., 45 km west of Hua Hin,
12.ix.1993, alici.
Description. - Fastigium verticis narrow, dorsally
furrowed; step-like declined to fastigium frontis.
Pronotum with disc flattened, especially in posterior
area, lateral angles rounded into paranota; transverse
sulcus V-shaped; with a faint indication of a medial
carinula; paranota longer than high (54:45), ventral
and posterior margins together broadly rounded;
humeral sinus distinct. Tegmen with radius sector
branching between 5.5 and 11.5 mm from base of
tegmen. Anterior femur with spines on both ventral
margins, on external side sometimes absent or only
one spine; meso- and postfemora with spines on ven-
tro-external margin; genicular lobes of all legs
bispinose. Anterior tibia with dorsal and ventral, ex-
ternal and internal apical spurs.
Male: Stridulatory file with circa 32-35 teeth
which are large and very widely spaced in circa basal
half, gradually decreasing in size in third quarter and
terminating in a series of minute teeth in apical quar-
ter (figs. 18-19). About 14-16 teeth may be regarded
as large and useful for sound production; the minute
teeth at apex are probably without function. Tenth
abdominal tergite slightly prolonged behind, apex
truncate; central area setose and with a distinct punc-
ture in middle. Epiproct long tongue-shaped; apex
subtruncate. Cerci curved, widest in basal area, slight-
ly narrowing in middle and slightly widening again
towards apex; apical area compressed and twisted-tri-
angular, very apex acute and curved cephalad (figs.
42-43). Subgenital plate narrow, only slightly curved,
split into two parallel lobes in apical quarter to apical
third (37:93 - 20:82) (figs. 63-64, 83).
Female: Tenth abdominal tergite short, transverse
with a distinct puncture in middle. Epiproct long
tongue-shaped, apex short-acutely projecting in mid-
dle. Cerci moderately curved, apex subacute (fig.
148). Subgenital plate long-triangular, apex short-
roundly excised (fig. 1 1 1). Gonangulum of ovipositor
with a large, semimembranous, ventro-apical ap-
pendage (figs. 120, 127).
Coloration: Green with or without a brown longi-
tudinal band on vertex, disc of pronotum and dorsal
area of tegmen (or part of these organs); compound
eyes brown; antenna concolorous or dorsal side of fla-
gellum infumate (antenna directed anteriorly), scapus
and pedicellus with or without some reddish pattern.
Tegmen with more or less distinct aggregations of
dark dots in the cells mainly in apical half of tegmen
and between radius and cubitus. Abdominal tergites
often reddish brown in middle and with or without
some reddish dots in lateral areas.
Measurements (length in mm): Body 6 22-23, 9
20-25; pronotum cT 4.2-4.5, 9 3.8-4.5; tegmen 6
33.0-35.0, 9 30.5-37.0; tegmen width 6 5.5 - 6.2,
9 5.5-6.0; anterior femur S 7.5-8.0, 9 7.5-8.0;
mesofemur â 9.5-10.5, 9 9.0-10.5; postfemur cT
21.0-23.0, 9 20.0-22.5; ovipositor 6.0-7.0.
Remarks. - The species was previously thought to
be widespread in the Oriental Region (Brunner 1878,
Karny 1926a, Jin & Xia 1994). However of the mate-
rial at hand, only specimens from Central and South-
ern Thailand, Malay Peninsula, Sumatra and Java can
be attributed to this species. The occurrence in other
regions has to be verified.
Stridulation (fig. 172). - The song of one male
from Lard Phrao (Bangkok, Thailand) was recorded
in the field during the night (temperature not mea-
sured but supposed to be about 25-28°C). Stridula-
tion consists of echemes of 2-3 crescendoing syllables.
The echemes can be rather regularly repeated for sev-
eral minutes. Syllable duration varied between 1 5-29
ms, echeme duration between 381-393 ms in the
two-syllabic and 1012-1318 in the three-syllabic
echemes. About ten single pulses can be recognised in
the syllables at a high time resolution. The song was
loudest at 10-15 kHz but higher frequencies were not
recorded with the field equipment. The sound of an
echeme can be circumscribed as "zip zip zip".
Elimaea (Elimaea) punctifera (Walker)
(figs. 26, 44, 61-62, 82, 160)
Phaneroptera punctifera Walker, 1869: 342. Holotype (â):
Bangladesh: Silhet (bmnh) [examined]. - Elimaea punc-
tiferaKirby 1906: 396; Karny 1926c: 23.
Redescription of holotype. - Fastigium verticis nar-
row, acute-angular in dorsal view with apex subacute,
dorsal surface furrowed. Pronotum with disc flattened
in posterior area, lateral margins rounded; transverse
sulcus slightly behind middle, V-shaped; paranota
longer than high (4:3); ventral and posterior margins
rounded together; humeral sinus distinct. Tegmen sur-
passing hind knees; radius sector branching circa 8 mm
from base. Legs I and II absent. Postfemur with spines
on ventro-external margin. Hind knees bispinose.
Male: Stridulatory file with about 53 teeth which
are large and spaced in basal half and gradually be-
coming narrower and denser towards apex (fig. 26).
Tenth abdominal tergite slightly prolonged and apex
subtruncate. Epiproct long tongue-shaped. Cerci
rather strongly curved, centre of curvature slightly be-
fore middle; apical area spatulate, rather long and
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Tijdschrift voor Entomologie, volume ui, 1998
step-like constricted against rest of cercus, with con-
vex external and concave internal surface, very apex
curved ventrad and subacute (fig. 44). Subgenital
plate narrow, rather strongly curved dorsad (arte-
fact?), with a weak medial carina, apex split in circa
apical quarter (figs. 61-62, 82).
Coloration: Rather uniformly green. Tegmen in
cells between media and cubitus and in apical half of
tegmen also in cells between radius and media with
aggregations of black dots; internal margin of tegmen
brownish.
Measurements of male (length in mm). Body 21;
pronotum 4.5; tegmen 36.0; tegmen width 6.0; post-
femur 27.0.
Remarks. — E. punctifera is a rather uniformly green
coloured species without striking characters. The
dark colour pattern caused by the black dots in the
cells of the tegmina are not as striking as described in
the key in Karny (1926a). Moreover, the subgenital
plate is not so strongly divided at the apex as sup-
posed by previous authors (Tinkham 1943, Bei-Bi-
enko 1955). Thus both authors probably misidenti-
fied other species as E. punctifera. Only the apical
quarter of the subgenital plate is divided which is the
same value as in E. Moris. The subgenital plate as well
as coloration which were previously thought to differ
between both taxa, are thus not distinctive. Both
species are very similar. They differ however strikingly
in the number of stridulatory teeth on the underside
of the left tegmen and less strikingly in the length of
the cerei and the shape of the apical area of the cerei.
E. punctifera was previously thought to be wide-
spread in the Oriental Region (Kirby 1906, Yin &
Xia 1994) and adventive to the Hawaiian fauna (He-
bard 1922b). However the material that I have seen
from Indochina and Hawaii is not conspecific with
the holotype. Thus the species is with certainty only
known from the type locality Silhet in Bangladesh.
Its' distribution is probably restricted to the Indian
subcontinent or to an even smaller range. The occur-
rence in other regions has to be verified.
Elimaea (Elimaea) thaii sp. n.
(figs. 24-25, 48, 57-58, 110, 129-130, 149-150,
156, 166, map 3)
Holotype 3 : Thailand, Tak prov. Mae Salid, Mon-
krating, 700m, 19.-21. v. 1988, S. Ingrisch, dab. -
Paratypes: Thailand: 1 9 , Tak prov. Mae Salid, Mon-
krating, 700m, 16.V.1988 (600-800m); IcT, same
data, 19.-20.ix. 1989; 23, same data, 11. -13.x. 1990;
1 ? , same data, 3 1 .vii. 1 992; 2 3 , same data, 1 .viii. 1 992
(700- 1000m); 1 3 , Tak prov., Doi Musoe, Agricultur-
al Research Station, 9.X.1990; \3, Tak prov.,
Umphang, 1 6.x. 1991; 1(5, 39, Chiang Mai prov.,
Doi Chiang Dao, 4.-7.vi.l986; 19, same data,
9.X.1991; ló\ 19, Chiang Mai prov., Phrao district,
Phrao - Ban Pradu, 26.-29.ix. 1985; 19, Chiang Mai
prov., lOkmNESamoeng, 10.x. 1991; 1 ó\ ^.Chi-
ang Mai prov., 5 km SE Samoeng, 4.vi.l997 ex larva;
11 3 , 59, same data but ex ovo; 1 9 , Chiang Mai
prov., Doi Suthep, 13-l400m, 8.X.1991; 29, same
data, 3.Ü.1997 ex larva; 19, Mae Hong Son prov.,
Samoeng - Pa Pae (Karen village - Kunsan Nai),
30.iv.1988; 1 9, Mae Hong Son prov., Pai - Soppong,
24.-261 1987, alici.
Description. - Fastigium verticis narrow, dorsally
furrowed, apex subacute. Pronotum with disc flat-
tened in posterior area, lateral angles rounded; trans-
verse sulcus short Y-shaped, slightly behind middle;
paranota about 1 . 1-1 .2x longer than high; ventral and
posterior margin rounded together; humeral sinus
distinct. Tegmen surpassing hind knees; radius sector
branching circa 7.5-12 mm from base.
Male: Stridulatory file with circa 34-42 teeth
which are large and somewhat spaced from each oth-
er in basal half and gradually becoming narrower and
denser towards apex (figs. 24-25). Tenth abdominal
tergite with apical margin broadly rounded; central
area setose and with a distinct puncture in middle.
Epiproct long tongue-shaped. Cerci rather short,
strongly curved; apical area short, compressed and
twisted, slightly curved, apex acute (fig. 48). Subgen-
ital plate narrow, split for slightly less than apical half
(apical four to five tenth) (figs. 57-58). Dorso-central
surface of medial phallus valves covered with warts.
Female: Tenth abdominal tergite with apex sub-
truncate or slightly concave; with a distinct puncture
in the middle. Epiproct long-triangular, apex subob-
tuse (figs. 149-150). Cerci moderately curved, slen-
der; apex varying from subacute to obtuse. Subgenital
plate long-triangular in general outline with a medial
furrow bordered by a carina at each side; apex sub-
truncate between two slightly projecting lateral angles
(fig. 110). Gonangulum of ovipositor with ventro-
apical appendage smaller than in E. subcarinata and
E. chloris (figs. 129-130).
Coloration: Male green; compound eyes brown;
antenna (stretched anteriorly) light brown on dorsal,
dark brown to black on ventral surface, with spaced
annulation; disc of pronotum with dark dots and
with a narrow brown medial band which is interrupt-
ed in middle by a fine white line. Tegmen with ag-
gregations of black dots in cells between media and
anal margin and in apical half of tegmen also between
radius and media. Abdominal tergites red in middle.
Variation: The brown band on pronotum some-
times indistinct or absent and leaving only the white
line and eventually a pair of irregular white lateral
lines present. The dark brown band may also extend
to the vertex and the dorsal area of the tegmen. In
86
Ingrlsch: Review of Elimaeini
darker individuals, the anterior and medial legs are of-
ten brownish or pale reddish brown and the anterior
margin of the tegmen reddish. The pronotum can be
with or without red dots and the anterior area of the
stridulatory field can be red.
Female varying from uniformly green (rarely
brown when alive) to very colourful. Head, antenna
and pronotum as in male. Pronotum with or without
dark brown or red dots; with or without a brown me-
dial band that is split by a fine white line (may be ex-
tended to vertex or not); with or without irregular
white lateral lines. Tegmen green with aggregations of
black dots in cells between radius and anal margin
varying from hardly expressed to very distinct, may
also be replaced by red dots; or cells between radius
sector and cubitus almost completely red and only the
veins green; or almost all of tegmen, body, and legs
suffused with red. Abdominal tergites usually red in
middle. Legs as in male; genicular region of anterior
leg (femur and tibia) often, but not always darkened.
Measurements (length in mm): Bodyc? 16-25, 9
20-25; pronotumc? 3.5-4.2, 9 3.5-4.2; tegmenc?
32.0-36.5, 9 31.5-37.0; tegmen width S 5.0-6.0, 9
5.0-6.0; anterior femur 3 7.5-9.0, 9 7.5-9.0; mesofe-
murc? 9.5-12.0, 9 9.5-11.0; postfemurc? 21.0-25.0,
9 20.0-24.5; ovipositor 6.0-6.5.
Remarks. - Part of the material of this species was
previously misinterpreted as belonging to E. punc-
tifera by Ingrisch (1990a). The stridulatory file and
the male cerei show however distinct differences to
the male holotype of E. punctifera. A large number of
specimens in the dab from numerous localities in
Thailand which were partly identified as E. chloris by
the late Dr. H.H. Karny probably also belong to this
species or to E. subcarinata; the stridulatory file was,
however, not checked.
The male subgenital plate of E. thaii is also similar
to that off. berezovskii Bei-Bienko, 1951, described
from the Sichuan province in China. Both species dif-
fer in the width of the tegmen which is 5.1-5.3 times
longer than wide in E. berezovskii (Bei-Bienko 1965),
but 6-7 times longer than wide in E. thaii. Other di-
agnostic characters are not well described for E. bere-
zovskii, especially the stridulatory file is unknown.
E. thaii is common in western and northern Thai-
land, where it can occur in the same habitat together
with E. subcarinata (map 3).
Etymology. - The name of this species refers to its
distribution.
Development. — Oviposition was in parenchyma of
leaves {Rubus). Egg development required about 42-
64 days at 18-25°C. There are six larval instars. The
first instar larvae are green; the longitudinal white
stripes are less conspicuous than in the young larvae
of E. subcarinata. Legs and antennae are yellowish
brown. When at rest, they sit as the adults with the
antennae and anterior legs stretched forwards and the
posterior legs spread in an acute angle from the body
(fig. 166). Several larvae became brown in subsequent
moultings, but almost all of them changed back to
green with the final ecdysis. Food plants were the
same as in E. subcarinata.
Stridulation (fig. 174). - Stridulation of one male
from Samoeng (Chiang Mai province, Thailand) was
recorded at 22.5°C. The male was kept with a female
larva and later with an adult female in the same cage.
On the latter occasion, the male female response
stridulation was also recorded.
Stridulation of the male kept together with the lar-
va consisted of single syllables which were repeated in
long and irregular intervals. In a 45 min continuous
recording, the male produced only 5 syllables. Ac-
cording to the higher number of stridulatory teeth,
syllable duration (122-213 ms) was much longer than
in both other Elimaea s. str. species.
On a later occasion, when a female answered the
males' syllable with a short and quiet click sound, the
male produced a second syllable. The female response
was 3042-3760 ms after the males' first syllable, while
the males' second syllable followed 6400-69 1 5 ms af-
ter the females' answer.
The main frequency of the male stridulation was at
8-18 kHz and thus rather loud for the human ear.
The sound of a syllable is the same as in the preceding
two species, but longer.
Discussion. - Male stridulation of the Elimaea and
Rhaebelimaea species so far studied is rather simple
and is composed of only a few short syllables. The dif-
ferences in stridulation of the three Elimaea species
however give evidence that despite of the great mor-
phological similarity they are three distinct species.
Moreover, the time pattern of the syllables they pro-
duce corresponds with the number of teeth on the
stridulatory file. This gives evidence that the stridula-
tory file is one of the most useful characters in Eli-
maeini (especially in Elimaea s. str.) and it should be
used for taxonomie purpose.
Elimaea (s. str.) nautica sp. n.
(figs. 27-28, 49-50, 55-56, 109, 145, map 3)
Holotype S : Thailand: Chanthaburi prov., Khao
Soi Dao, 29.-3 l.v. 1990, A. Lewvanich, V. Koon-
tong, S. Wangsuk, dab (Lot 4166). - Paratypes: 1 â ,
Thailand, Chanthaburi prov., Khao Soi Dao,
15.X.1985, ci. - usa: la, 1$, Hawaii, Big Island,
near Kaumana Cave, 7.viii.l993, ci.
Description. - Fastigium verticis narrow, dorsal
surface sulcate, in lateral view sinuate; step-like de-
clined to fastigium frontis. Pronotum with disc flat-
tened but lateral angles rounded; transverse sulcus V-
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Tijdschrift voor Entomologie, volume hi, 199s
or short Y-shaped; with a short and weak medial cari-
na before apex; anterior margin concave, posterior
margin rounded; paranota l.lx longer than high,
ventro-posterior angle rounded; humeral sinus dis-
tinct. Anterior femur with spines on both ventral
margins (on internal side more than on external);
meso- and postfemur with spines on ventro-external
margin. Tibial tympana conchate on both sides. An-
terior tibia with dorsal and ventral, external and in-
ternal apical spurs; dorsal surface sulcate.
Male: Stridulatory file with circa 28-29 teeth
which are large and widely spaced from each other
(figs. 27-28). Tenth abdominal tergi te with apical
margin widely rounded; central area setose and pro-
vided with a puncture in centre. Epiproct tongue-
shaped, apex broadly rounded. Cerci strongly curved,
rather short; apical area compressed and excavated on
cranial side; tapering towards acute apex (figs. 49-50).
Subgenital plate long and narrow, with a medial car-
inula in basal half; split into 2 subparallel lobes in cir-
ca apical half or more (apical five to six tenth); apical
lobes setose (figs. 55-56).
Female: Epiproct tongue-shaped. Cerci short,
slightly curved, stout in basal area, narrowing in
about middle, but then cylindrical and of subequal
width to apex; apex obtuse (fig. 145). Subgenital plate
long-triangular, apex rather broadly excised in mid-
dle, subtruncate between two short obtuse lateral pro-
jections (fig. 109).
Coloration: Green. Vertex and disc of pronotum
with a faint brown medial band split in middle by a
whitish line and with scattered brown dots, lateral an-
gles with a little distinct white line. Tegmen green,
anterior margin reddish or anterior area indistinctly
infumate, with aggregations of black dots mainly be-
tween radius and anal margin.
Variation. In the female at hand, paranota and ab-
domen with red dots; medial band on disc of prono-
tum reddish, not split in middle.
Measurements (length in mm): Body S 21-22, 9
21; pronotum cT 4.2-4.5, 9 4.8; tegmen 6 36.5-
37.5, 9 40.0; tegmen width S 5.5-63, 9 6.0; ante-
rior femur S 9.0, 9 10.0; mesofemur S 11.5-12.0,
9 12.0; postfemur Ó* 23.0-24.0, 9 24.0; ovipositor
8.0.
Remarks. - This species agrees with the description
of E. punctifera by Bei-Bienko (1955), not Walker
(1869). A re-investigation of Walkers' type shows
however that both species are quite different.
Material of E. nautica at hand was collected in
south-eastern central Thailand (map 3). It is possible
that its range spread from there to Cambodia and
South Vietnam. E. nautica is also adventive to the
Hawaiian fauna. Whether other Elimaea species, e.g.
E. punctifera as recorded by Hebard (1922b), also oc-
cur on the Hawaiian Islands was not investigated.
Etymology. — The name of this species is derived
from Latin "nauticus" = seaman. It refers to its ability
to cross the ocean and become an adventive species in
Hawaii.
Elimaea poaefolia— group
Type species: Locusta (Phaneroptera) poaefolia De Haan,
1842; here designated.
Diagnosis. - Elimaeini with anterior femora curved
as in phasmids. Tibial tympana covered by a conchate
fold on both sides. Width of tegmen in middle slight-
ly narrower than length of pronotum in male or
width subequal to length of pronotum in female; in
male tegmen often widening towards apex (for about
1 mm); radius sector branching distinctly before mid-
dle of tegmen. Phallus membranous. Ovipositor
elongo-falcate, margins serrulate near apex; gonangu-
lum with ventro-apical angle projecting.
Remarks. - There are two females from northern
Thailand at hand that cannot be assigned to any de-
scribed species and that obviously represent two dif-
ferent species. As the corresponding males are not
known, I hesitate to name the new taxa, but an infor-
mal description of the two females is given below.
Distribution. - The poaefolia- group was previous-
ly known with one species each from Java, Sumatra,
Borneo and Malaysia. The material at hand proves
that the range of the group extends at least to the
mountains of northern Thailand (maps 1,3).
Elimaea (Elimaea) sp. 1
(figs. 115, 125, 138, map 3)
Material studied. - Thailand: 1 9 , Chiang Mai prov.,
Chiang Dao, iv.1958, Phon, dab (Lot 2409).
Description. - Female: Fastigium verticis narrow,
sulcate, apex obtuse and step-like declined to
fastigium frontis. Pronotum with disc nearly flat and
with a low medial carina which is replaced by a fur-
row behind transverse sulcus and distinct again before
apex; transverse sulcus arcuate; paranota circa 1.5
times longer than high with a longitudinal carina be-
low middle of height, ventral margin slightly convex,
anterior and posterior angles rounded, humeral sinus
distinct. Tegmen narrow, subparallel-sided, apex
rounded; radius sector branching distinctly before
middle. Anterior femur with spines on ventro-inter-
nal margin. Mesofemur with spines on ventro-exter-
nal margin; postfemora lacking. Knee lobes of pro-
and mesofemur bispinose. Anterior tibia with dorsal
and ventral, external and internal apical spurs. Ante-
rior and medial legs very thin.
Male unknown.
Female: Tenth abdominal tergite with apical mar-
88
Ingrisch: Review of Elimaeini
gin subsinuate. Epiproct long-triangular with apex
obtuse; basal half bulging and with a medial furrow,
apical half flattened. Cerci long-conical, moderately
curved, apex subacute. Subgenital plate transverse,
with a medio-longitudinal fold (artefact?); apical mar-
gin transversely truncate but slightly excised in mid-
dle; apico-lateral angles with two faint, obtuse,
bulging swellings (fig. 115). Gonangulum of oviposi-
tor with a strong projection pointing (ventro-)apicad
(figs. 125, 138).
Coloration: Uniformly yellowish brown (dis-
coloured?). Pronotum with dark dots on disc and lat-
eral angles. Tegmen with light veins and veinlets and
dark cells, less expressed in anterior-apical area.
Measurements of female (length in mm): Body 23;
pronotum 4.5; tegmen 36.0; tegmen width 4.5;
mesofemur 13.5; ovipositor 8.3.
Remarks. - This species is close to the following, it
differs by the transverse subgenital plate, the stouter
process of the gonangulum which is pointing apicad
not ventrad and the lateral lobes of pronotum with
the ventral margin broadly rounded not truncate.
Elimaea (Elimaea) sp. 2
(figs. 114, 126, 137, 139, map 3)
Material studied. - Thailand: 1 5 , Tak prov., Mae Salid,
Monkrating, 700m, 1 .viii. 1992, at night, ci.
Description. - Fastigium verticis narrow, sulcate,
apex obtuse and step-like declined to fastigium fron-
ds. Pronotum with disc nearly flat and with a medial
carina which becomes subobsolete in metazona, later-
al angles rounded; transverse sulcus broad-Y-shaped;
anterior margin concave, posterior margin rounded;
paranota circa 1.5 times longer than high with a lon-
gitudinal carina in about middle of height, ventral
margin substraight, anterior and posterior angles
rounded, humeral sinus present. Tegmen surpassing
hind knees, narrow, parallel-sided, apex rounded; ra-
dius sector branching distinctly before middle of
tegmen. Anterior femur with spines on ventro-inter-
nal margin, meso- and postfemur with spines on ven-
tro-external margin. Knees of all femora bispinose.
Anterior tibia with dorsal and ventral, external and
internal, apical spurs.
Male unknown.
Female: Tenth abdominal tergite with apical mar-
gin subsinuate. Epiproct subpaiallel-sided in circa
basal half, with a medial furrow and lateral bulges, flat
and long-triangular in apical half. Cerci long-conical,
moderately curved, apex subobtuse (fig. 139). Sub-
genital plate with disc circa pentagonal in general out-
line and with a medial furrow in apical half; lateral
area sloping and prolonged into spinose projections
pointing apicad (fig. 1 14). Gonangulum of ovipositor
with a short, conical, obtuse projection pointing ven-
trad (figs. 126, 137).
Coloration: Green; pronotum and tergites with
black dots. Tegmen with black dots forming three in-
distinct bands in radial and medial areas; costal area
with red spots.
Measurements of female (length in mm): Body 22;
pronotum 4.0; tegmen 34.0; tegmen width 4.0; post-
femur 24.5; ovipositor 8.0.
Acknowledgements
I would like to thank Drs. J. van Tol (Leiden), Dr.
Yayuk R. Suhardyono (Bogor) and Mrs. Boopa Laos-
inchai (Bangkok) for their much appreciated help
during my visits in the following museums: Nationaal
Natuurhistorisch Museum Leiden, Museum Zoolog-
icum Bogoriense, Department of Agriculture, respec-
tively. For the loan of material I am indebted to the
following persons: Dr. A. Kaltenbach and Dr. U. As-
pöck (Vienna); Dr. D. R. Ragge (London), Dr. R.
Poggi (Genoa), Dr. T. Kronestedt (Stockholm), and
Dr. M. S. Shishodia (Calcutta). Dr. F. Willemse
(Eygelshoven) helped me with tracing references. My
visits at the museums in Leiden and Bogor were sup-
ported by the German research council (DFG, In 2-
2). Thanks go also to Dr. D. C. F. Rentz (Canberra)
and Dr. E. J. van Nieukerken for their valuable com-
ments on an earlier version of the manuscript.
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Bei-Bienko, G.Y., 1955. Investigations on the fauna and sys-
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Bei-Bienko, G.Y., 1962. Results of the Chinese-Soviet zoo-
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Bei-Bienko, G.Y., 1965. Tettigonioidea Phaneropterinae.
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Bolivar, L, 1914. Examen de un pequeno lote de Ortópteros
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161 (Chinese with English summary).
Received: 21 October 1997
Accepted: 11 March 1998
90
Ingrisch: Review of Elimaeini
Abbreviations for species names in figs. 2-150:
api E. (R.) apicata sp. n.
app E. (E.) appendiculata Brunner, 1 878
chi E. (E.) Moris (De Haan, 1842)
cue E. (H.) cuculiata Ingrisch, 1990
cur E. (R.) curvicercata (Brunner, 1891)
heb E. (R.)hebardi Karny, 1926
kra E. (R.) kr aussi Karny, 1926
E. (O.) leeuwenii Ka.my, 1926
E. (R.) maninjauensis sp. n.
E. (R.) mentaweii sp. n.
E. (O.) minor (Brunner, 1891)
E. (R.) modiglianii sp. n.
E. (E.) nautica sp. n.
E. (R.) neglecta Karny, 1 926
E. (R.) parumpunctata (Serville, 1839)
pen E. (R.) pentaspina sp. n.
pro E. (H.) procera Ingrisch, 1990
pse E. (R.) pseudochlorissp. n.
pun E. (E.) punctifera (Walker, 1869)
roa E. (R.) roseoalata (Brunner, 1891)
sia E. (R.) siamensis Karny, 1926
sig E. (R.) signata (Brunner, 1 878)
sin E. (R.) sinuata sp. n.
spi E. (E.) sp. 1
sp2 E. (E.) sp. 2
sub E. (E.) subcarinata (Stai, 1861)
sum E. (R) sumatrana Ka.my, 1926
tha E. (E.) thaii sp. n.
tra E. (R.) transversa Ingrisch, 1990
wil E. (R.) willemsei Karny, 1 926
H specimen from Hawaii
T specimen from Thailand.
91
Tijdschrift voor Entomologie, volume ui, 1998
Figs. 2-17. -2-16, Stridulatory files of Elimaea species: 2, E. (O.) leeuwenii; 3, E. (O.) minor-A, E. (R.) apicata;5, E. (R.) sin-
uata; 6, E. (R.) parumpunctata; 7, E. (R.) mentaweii; 8, E. (R.) maninjauensis; 9, E. (R.) pseudochloris; 10, E. (R.) kraussi; 11,
E. (R.) curvicercata; 12-13, E. (R.) modiglianii (13 oblique profile); 14-15, E. (H.) procera (1 5 oblique profile); 16, E. (H.) cu-
culiata; 17, dorsal field of right tegmen of £ (R.) kraussi.
92
Ingrisch: Review of Elimaeini
1^])')'}}3}}33'jJJiJJ>JîU:n//;/,//\
25
g 1 0 3 3 ì 3 3 1 J 1 1 11 HjiüllillimiJIJUJlin/iM,,
26
O OOOOQOOOGOOOOOOoaoa///,^
3 0 0 Tj£iW£j »» nmnÎÏJî;.
27
28
Figs. 18-28. Stridulatory files of Elimaea species. - 18-19, E. (E.) Moris (18, Java, syntype; 19, Bangkok, Thailand); 20-23,
E. (E.) subcarinata (20, Hongkong, syntype; 21, Indochina, type of E. (E.) appendiculata; 22-23, Doi Khun Tan, Thailand);
24-25, E. (E.) thaii (24, Monkrating, Thailand; 25, Doi Musoe, Thailand); 26, E. (E.) punctifera, Silhet, Bangladesh, type;
27-28, E. (E.) nautica (27, Khao Soi Dao, Thailand; 28, Caumana Cave, Big Island, Hawaii).
93
Tijdschrift voor Entomologie, volume i4i, 1998
Figs. 29-41. Male cerei of Elimaea species (dorsal view; A = apex in ventral view). - 29, E. (R.) willemsei; 30, E. (R.) sinuata;
31, E. (R) modiglianii; 32, E. (R.) parumpunctata; 33, E. (R) curvicercata; 34, E. (R.) kraussi; 35, E. (R) sumatrana; 36, E.
(R.) pseudochloris; 37, E. (R.) apicata; 38, E. (R.) signata; 39, E. (R) maninjauensis; 40, £. (R.) mentaweii; Al, E. (R.) hebar-
di. [35, 41 after Hebard 1922; 38 after Karny 1926a; 29 after Karny 1926b; all modified].
94
Ingrisch: Review ofElimaeini
Figs. 42-54. Male cerei of Elimaea species (dorsal view, A = apex in ventral view). - 42-43, E. (E.) chlorìs (42, Java; 43,
Bangkok, Thailand); 44, E. (E.) punctifera, Silhet, Bangladesh, type; 45-47, E. (E.) subcarinata (45, Hongkong, syntype; 46,
Indochina, type of E. (E.) appendiculata; Al, Doi Khun Tan, Thailand); 48, E. (E.) thaii, Doi Musoe, Thailand; 49-50, E.
(E.) nautica (49, Caumana Cave, Big Island, Hawaii; 50, Khao Soi Dao, Thailand); 5 1 , E. (O.) minor, Tretes, Java; 52, E. (O.)
leeuwenii, Thong Pha Phum, Thailand; 53, E. (H.) procera, Khao Soi Dao, Thailand; 54, E. (H.) cuculiata, Erawan, Thailand.
95
Tijdschrift voor Entomologie, volume
Figs. 55-83. Male subgenital plate of Elimaea species in ventral view (82-83 in lateral view). - 55-56, E. (E) nautica (55 Cau-
mana Cave, Big Island, Hawaii; 56, Khao Soi Dao, Thailand); 57-58, E (E.) thaii (57, Doi Musoe; 58, Monkrating, Thai-
land); 59-60, E (E.) subcannata, Doi Khun Tan, Thailand (59, air-dried; 60, freeze-dried); 61-62, E. (E.) punctifera, type
(61, ventro-apical view; 62 ventral view); 63-64, E (E.) Moris (63, Bangkok, Thailand; 64, Palabuan Ratu, Java); 65 E (E)
^cannata Hongkong, syntype; 66, E (R) curvicercata; 67, E (R) kraussi; 68, E (R.) parumpunctata; 69, E (R) sumatrana;
70 E (RJroseoalata; 71, E (R.) sinuata; 72, E (R.) modiglianü; 73, E (R.) pseudochloris; 74, E (R.) apicata; 75, E (R)
hebardi; 76, E (R) mamnjauensis; 77, E. (R) mentaweii; 78, E (O.) leeuwenü; 79, E (O.) minor; 80, E (H.) cuculiataci,
E (H) procera; 82, E (E.) punctifera, type; 83, E (E) chloris, Java. [69, 75 after Hebard 1922; 70 after Karny 1926c; 78, 81
after Ingnsch 1990a; all modified].
96
Ingrisch: Review of Elimaeini
Figs. 84-96. - 84-86, Elimaea (R.) mentaweii. 84, abdominal apex; 84a, epiproct; 85, phallus sclerite in medial view; 86, same
in lateral view. - 87-96, phallus sclerites of Elimaea species in lateral view: 87, E. (R.) parumpunctata; 88, E. (R.) curvicercata;
89, E. (R.) modiglianii; 90, E. (R.) sinuata (paired dorsal and unpaired ventral sclerites); 91, E. (R.) apicata; 92, E. (R.) kraus-
si; 93, E. (R.) pseudochloris; 94, E. (R.) maninjauensis; 95, E. (H.) procera; 96, E. (H.) cuculiata. - Abbreviations: tt tenth ab-
dominal tergite, e epiproct, c cercus, ph phallus, sc phallus sclerite, sg subgenital plate.
97
Tijdschrift voor Entomologie, volume i4i, 1998
Figs. 97-1 18. Female subgenital plate of Elimaea species. - 97, E. (R.) parumpunctata; 98, E. (R.) transversa; 99, E. (R.) man-
injauensis; 100, E. (R.) mentaweii; 101, E. (R.) pentaspina; 102, E. (R) modiglianii; 103, E. (R.) roseoalata; 104, E. (R.) hebar-
di; 105, E. (R.) kraussi; 106, E. (R.) neglecta; 107, E. (R) siamensis; 108, E. (R.) signata; 109, E. (E.) nautica, Caumana Cave,
Big Island, Hawaii; 1 10, E. (E.) thaii, Doi Chiang Dao, Thailand; 1 1 1, £ (E.) chloris, Bogor, Java ; 112-113, E. (E.) subcar-
inata (112, Hongkong, syntype; 113, Doi Khun Tan, Thailand, freeze-dried); 114, E. (E.) sp. 2; 115, E. (E.J sp. 1; 116, E.
(O.) leeuwenii; 1 17, E. (H.) cuculiata; 1 18, E. (H.) procera; [103, 105 after Karny 1926a; 104 after Karny 1926b; 106-108 af-
ter Karny 1926c; all modified].
98
Ingrisch: Review ofElimaeini
Figs. 119-126. Ovipositor of Elimaea species. - 119, E. (O.) leeuwenii; 120, E. (E.) chloris, Bogor, Java; 121, E. (R.)
parumpunctata; 122, E (R.) maninjauensis; 123, E. (R.) mentaweii; 124, E (R.) pentaspina; 125, E. (EJsp. 1; 126, E. (EJsp.
2 - Abbreviations: ap appendage, c cercus, dv dorsal valve, e epiproct, g gonangulum, sg subgenital plate, w ventral valve.
99
Tijdschrift voor Entomologie, volume ui, 1998
Figs. 127-150. - 127-138, gonangulum at base of ovipositor of Elimaea species: 127, E. (E.) chloris, Bogor, Java; 128, E. (E.)
subcarinata, Doi Khun Tan, Thailand; 129-130, E. (E.) thaii, (129, Doi Chiang Dao; 130, Monkrating, Thailand); 131, E.
(E.) nautica, CaumanaCave, Big Island, Hawaii; 132, E. (R.) modiglianii; 133, E. (R.) transversa; 134, E. (R.) pentaspina; 135,
E. (R.) mentaweü; 136, E. (R) parumpunctata; 137, E. (E.) sç. 2; 138, E. (E.) sp. 1 - 139-150, left female cercus of Elimaea
and Rhaebelimaea species: 139, E. (E.)sp. 2; 140, E. (R) modiglianii; 141, E. (R.) transversa; 142, E. (R) pentaspina; 143, E.
(R) maninjauensis; 144, E. (R.) mentaweü; 145, E. (E.) nautica, Caumana Cave, Big Island, Hawaii; 146-147, E. (E.) sub-
carinata (146, Doi Khun Tan, Thailand; 147, Hongkong, syntype); 148, E. (E.) chloris, Bogor, Java; 149-150, E. (E.) thaii
(149, Doi Chiang Dao; 150, Monkrating, Thailand). - Abbreviations: ap appendage of ventral ovipositor valve, dv dorsal
ovipositor valve, w ventral valve.
100
Ingrisch: Review of Elimaeini
[Figs. 151-156 see page 102]
Figs. 157-163. Habitus of Elimaea and Rhaebelimaea species. - 157, E. (E.) chloris, syntype male (rmnh); 158, E. (E.) sub-
carinata, syntype male (nrs); 159, E. (R.) mentaweii, holotype male (mzb); 160, E. (E.) punctifera, holotype male (bmnh);
161, E. (R.) pseudochloris, holotype male (dab); 162, E. (R.) sinuata, holotype male (mzb); 163, E. (E.) appendiculata, holo-
type male (nhmw).
101
Tijdschrift voor Entomologie, volume ui, 1998
Figs. 151-156. Habitus of Elimaea species. - 151, E. (O.) minor, male, Tretes, Java; 152, E. (R.) parumpunctata, female, Gu-
nung Salak, Java; 153, E. (R.) maninjauensis, holotype male, Lake Maninjau, West Sumatta; 154, E. (R.) apicata, holotype
male, Khao Sok, Thailand; 155, E. (R.) curvicercatct, Tretes, Java; 156, E. (E.) thaii, paratype male, Samoeng, Thailand
[Figs. 157-163: seepage 101] &
102
Ingrisch: Review ofElimaeini
Figs. 164-168. - 164, Elimaea (E.) subcarinata, female with spermatophore. - 165-168, first instar larvae of: 165, E. (E.) sub-
carinata; 166, E. (E.) thaii; 167, E. (R.) parumpunctata; 168, Ectadia fiilva.
103
Tijdschrift voor Entomologie, volume ui, 1998
169) Êctadia fulva; Doi Suthep
170) Elimaea (Hemielimaea) cuculiata; Erawan
171) Elimaea (Rhaebelimaea) parumpunctata; Gunung Salak
24.5°C, short song
27°C, long song
Hl
1
il
Is
'M
1 s
24.5°C, short song
-- .uu.mil
'Wir
il
"•"•"'Wfffffl
w
'1 ' Ml|
10 ms
27°C, long song
k
À A À
4
A
f
TT*
1
100 ms
Figs. 169-171. Oscillograms of stridulation. - 169, Ectadia fulva: upper row, sequence of echemes; lower row, echeme on ex-
panded time scale; 170, Elimaea (H.) cuculiata: upper row: series of three echemes; middle row: echeme on expanded time
scale; lower row, last three syllables of an echeme; 171, Elimaea (R.) parumpunctata: upper row left, one-syllabic song; upper
row right, multi-syllabic song; middle row, syllable of short song on expanded time scale; lower row, multi-syllabic song on
expanded time scale.
104
Ingrisch: Review of Elimaeini
172) Elimaea (Elimaea) chlor is; Bangkok, Lard Phrao
1
I
'
1 s
nil
"'f
1 10 ms
173) Elimaea (Elimaea) subcarinata; Doi Khun Tan
174)
Elimaea (Elimaea) thaii; Samoeng
22.5°C
'
ls
10 ms
9
i'
r
500 ms
Figs. 172-174. Oscillograms of stridulation. - 172, Elimaea (E.) chloris: upper row, series of echemes; lower row, syllable on
expanded time scale; 173, Elimaea (E.) subcarinata: upper row, series of three syllables; lower row, syllable on expanded time
scale; 174, Elimaea (E.) thaii: upper row, one-syllabic song; middle row, syllable on expanded time scale; lower row, male-fe-
male response song.
105
Tijdschrift voor Entomologie, volume hi, i998
Elimaea s. str.
poaefol /«-group
Hemielimaea
Orthelimaea
Ectadia
Map 1 . Known areas of the genera and subgenera of Elimaeini, without subgenus Rhaebelimaea.
106
Ingrisch: Review of Elimaeini
°s
10
95
100
105
110
115
120
125 °E
Map 2. Distribution of Elimaea (Rhaebelimaea) species in South East Asia. Species without precise locality data are tentative-
ly placed but not encircled. 1 parumpunctata, 2 kraussi, 3 curvicercata, 4 hebardi, 5 willemsei, 6 mentaweii, 7 sinuata, 8 man-
injauensis, 9 sumatrana, 10 modiglianii, 11 roseoalata, 12 adspersa, 13 signata, 14 spinigera, 15 neglecta, 16 siamensis, 17
pseudochloris, 18 apicata, 19 transversa, 20 pentaspina, 21 malayica, 22 caricifolia, 23 moultonii, 24 lamellipes, 25 longicercata,
26 puncticosta, 27 bakeri, 28 filicauda.
107
Tijdschrift voor Entomologie, volume mi, 1998
Map 3. Distribution of the species of Elimaea and Ectadia in Thailand; without subgenus Rhaebelimaea.
108
T. RUITEN' & O. KARSHOLT2
'embl, Heidelberg, Germany
Zoologisk Museum, Copenhagen, Denmark
BRYOTROPHA MUNDELLA (DOUGLAS): A NEW
SYNONYM OF BRYOTROPHA UMBROSELLA (ZELLER)
(LEPIDOPTERA, GELECHIIDAE)
Rutten, T. & O. Karsholt, 1998. Bryotropha mundella (Douglas): a new synonym of Bryotropha
umbrosella (Zeiler) (Lepidoptera Gelechiidae). - Tijdschrift voor Entomologie l4l: 109-114,
figs. 1-13.[issn 0040-7496]. Published 30 November 1998.
The status of Bryotropha mundella (Douglas) is discussed. Its genitalia are found to be identical
to those of B. umbrosella (Zeiler). As to the forewings of these two taxa, numerous intermedi-
ates exist spanning the whole range between the whitish nominate form of B. mundella and the
blackish nominate form of B. umbrosella. Geographically, B. mundella co-distributes with B.
umbrosella. The two moths are found in the same period of the year and are active during the
same time of the day. We therefore conclude that B. mundella is a light form of B. umbrosella.
Lectotypes of Gelechia umbrosella Zeiler, 1839, and of its synonyms G. munde ila Douglas, 1850
(syn. n.), G portlandicella Richardson, 1890 and G anacampsoidella Hering, 1924 are desig-
nated. [B. umbrosella is recorded from Spain for the first time].
Correspondence: T. Rutten, embl, Meyerhofstrasse 1, D-69012 Heidelberg, Germany. E-mail
rutten@embl-heidelberg.de
Key words. - Lepidoptera; Gelechiidae; new synonym.
The holarctic genus Bryotropha Heinemann, 1870
consists of almost 80 nominal species, about 40 of
which are currently recognised as valid. The genus
has never been revised, and especially in the south-
western part of the palaearctic region there are a num-
ber of undescribed species. Even among specialists,
Bryotropha is regarded as one of the more difficult
genera of Microlepidoptera.
Pierce and Daltry (1938) divided the genus into two
subgenera, Bryotropha s. str. and Mniophaga Pierce &
Daltry, 1938. In the subgenus Bryotropha the male
genitalia have a strong, specialised gnathos while the fe-
male genitalia have a plate-like signum with two trans-
verse ridges (Sattler 1971). The subgenus Mniophaga
was reserved for species in which the male has a rather
small gnathos and the female a plate-like signum with
strong spikes at the corners. There are, however, inter-
mediate taxa, and Sattler (loc. cit.) concluded that a
subdivision of Bryotropha is unjustified. Differences in
the genitalia are distinct between members of the first
group ('subgenus' Bryotropha), but less distinct be-
tween species of the second group ('subgenus' Mnio-
phaga) (Rutten, in press). The latter include the small,
'dark' Bryotropha species of northern and central Eu-
rope, which often cause much confusion. The light
coloured B. mundella (Douglas) also belongs to this
group and this moth is the subject of the present paper.
According to the original description by Douglas
(1850), the distinct feature of B. mundella is its light
greyish ground colour which distinguish this moth
from other members of the genus. Over the years,
however, the name B. mundella has been given to a
whole range of small Bryotropha with ground colour
varying from nearly white to dark grey. Doubts were
raised whether all these forms are representatives of B.
mundella (see e.g. Pierce & Daltry 1938).
Unfortunately, individual variations within Bry-
otropha species are poorly investigated and the geni-
talia of B. mundella have never been properly described.
The drawings published by Pierce and Metcalfe (1935)
are not suited for an identification. In the best study
on Bryotropha so far, Svensson (1962) describes the
genitalia of B. mundella as resembling those of B. um-
brosella (Zeiler). The description and drawings of B.
mundella given in Piskunov (1990: 970, 977) proba-
bly refer to B. afftnis or B. similis (Stainton). Speci-
mens identified by Piskunov as B. mundella and ex-
amined by us belonged to B. similis.
B. mundella is a rather scarce moth, especially on
mainland Europe. Only a handful were known from
109
Tijdschrift voor Entomologie, volume i4i, 1998
I
Figs. 1-8. Adults of Bryotropha. - 1, nominate fotm of B. umbrosella; The Netherlands, Zwanenwatet; 2, fotewing of B. um-
brosella (enlargement of fig. 1); 3, forewing of B. umbrosella with a distinctive lighter ground colour; The Netherlands, Ter-
schelling; 4, forewing of B. umbrosella revealing strong irroration with whitish scales; The Netherlands, Ameland; 5, extreme
whitish form of B. mundella with distinct stigmata and dark scales limited to the termen; Ireland, Fanore, Co. Claire; 6,
forewing of B. mundella showing traces of darker scales near the base of the wing; The Netherlands, Zwanenwater; 7, forewing
of B. mundella with predominance of greyish scales; The Netherlands, Terschelling; 8, forewing of a very dark B. mundella
with clearly defined costal and tornai patches; Ireland, Mannin Bay, Co. Galway.
110
Ruiten & Karsholt: Bryotropha umbrosella
mjf
10
Figs. 9-12. Adults of Bryotropha. -9, lectotype of B. umbrosella; 10, leccotype of B. mundella; 1 1, nominate form of B. affi-
nisi Great Britain, Norwich, Norfolk, 9. IV. 1995; 12, light coastal form of B. affinisi The Netherlands, Zwanenwater.
The Netherlands until the early 1 990's, when a small se-
ries were collected from the Frisian islands (Küchlein
1993). According to the genitalia, however, the moths
involved belonged to two other species; specimens with
a yellowish to brownish colour belonged to B. affinis
while those with a whitish or greyish colour, typical for
B. mundella, invariably had genitalia similar to B. um-
brosella (Zeiler). The implications were obvious, for, if
specimens with the external features of B. mundella have
genitalia which are identical to those of B. umbrosella,
one may ask whether B. mundella is indeed a separate
species and not just a light form of B. umbrosella.
Material and methods
Female genitalia were mounted in a ventro-dorsal
position. In the case of the male genitalia, lateral
mounting was preferred to ventro-dorsal mounting
since the latter procedure can distort the shape of the
gnathos, which is an essential characteristic in Bry-
otropha. Additional slides of male genitalia were made
using the unrolling technique.
Type material is preserved in the collections of the
Dorset County Museum (dorc), Dorchester, UK;
The Natural History Museum (bmnh), London, UK;
National Museum of Natural History (rmnh), Lei-
den, The Netherlands; Zoologisk Museum, Universi-
ty of Copenhagen (zmuc), Copenhagen, Denmark,
and in the Zoological Museum, University of Helsin-
ki (mzhf), Helsinki, Finland.
Results and discussion
Bryotropha umbrosella (Zeiler)
Gelechia umbrosella Zeiler, 1839: 201. Lectotype 9: [Po-
land; Glogów.] 'Gross Glogau, Silesia. 6.vi.l834, Zeiler
Coll'.; 'Walsingham Collection [B.M.] 1910-427'; 'Ge-
lechia umbrosella Zell. Isis p. 201 (1839) Type [male]; ab-
domen missing'; 'Lectotype [female] Gelechia umbrosella
Z. teste K. Sattler, 1961' (bmnh). [examined]
Gelechia mundella Douglas, 1850: 64. Lectotype S: [Great
Britain; New Brighton, Cheshire] '17.52'; 'England, Dgl
coll. (Mason 1906)'; 'Walsingham collection [B.M.]
1910-427'; 'Gelechia mundella Dgl. Tr. Ent. Soc. Lond.
(2)1, p. 64 (1850) Type [male]'; 'Genitalia no. 139'.
(bmnh) [examined] syn. n.
Gelechia portlandicella Richardson, 1890: 29. Lectotype 9:
[Great Britain; Portland, Dorset.} 'Lectotype Gelechia port-
landicella Richardson'; 'Portland, Dorset, Richardson Col-
lection 1889'; 'teste T. Rutten 1998' (dorc). [examined]
Bryotropha umbrosella [race] fidvipalpellajoasmis, 1908: 793.
Gelechia anacampsoidella Hering, 1924: 80. Lectotype â :
[Finland] 'Tvärminne'; '15.6.1921'; 'Kanerva'; 'Type;
Gelechia anacampsoidella, det. Mart. Hering m[ale]';
'Mus. Zool. Höfors; Spec. typ. No.7013; Gelechia ana-
campsoidella Her'.; [genitalia in glycerine vial]; 'Lectotype
Gelechia anacampsoidella Hering des. O. Karsholt 1998'
(mzhf) [examined].
Bryotropha oppositella (Thunberg, 1794) sensu Benander
1961: 245 [misidentification].
Bryotropha fidiginosella Snellen, 1882 sensu Lempke 1976:
25 [misidentification].
Ill
Tijdschrift voor Entomologie, volume i4i, 1998
Diagnosis
Adult. - Wingspan 9-12 mm. Head with very dark
greyish brown vertex and yellow to ochreous frons.
Thorax as forewing. Forewing very dark greyish brown
with blackish discal and plical stigmata barely visible;
plical stigma often with a few conspicuous white scales
beyond; costal and tornai patches white and usually
very prominent, rarely fused to form a fascia.
Variation. - In the form mundella the forewing is
irrorate with whitish or greyish scales. This makes the
blackish stigmata more prominent but obscures the
whitish tornai and costal patches. In extreme cases the
tornai and costal patches are no longer visible.
Male genitalia. - Gnathos slender, in a strong
(2=120°) but gradual curve, terminating in acute apex.
Tegumen alongside gnathos covered with 5-30 small
spines. Aedaegus long and slender, with bulbous base
and 'whip-like' apex.
Female genitalia. - Posterior margin of segment 8
with a median extension along dorsum, ventral part
of segment 8 densely covered with strong wedge-
shaped microtrichia. Lamella postvaginalis broad and
without lobes; signum square to rectangular with
strong spines at the corners.
Remark. - The genitalia of B. umbrosella and relat-
ed species are figured by Rutten (in press) .
Biology
The larva feeds until May in a spun tube among
mosses (Schütze 1931: 13). Imago univoltine from
late May to early August.
B. umbrosella frequents dry sandy places both in-
land and in coastal areas. B. mundella is always found
on locations also inhabited by B. umbrosella. The two
moths appear over the same period of the year. Field-
work by the second author in Jutland, Denmark,
found B. mundella flying amongst true B. umbrosella.
Both were active just before sunset. In contrast, B.
similis, which also occurred on that locality, was on
the wing between sunset and dark.
Distribution
B. umbrosella is widespread in northern and central
Europe; less common in southern Europe; absent
from Italy and Greece while records for Portugal need
confirmation (Karsholt & Riedl 1996: 109). A male
specimen identified during our study becomes the
first record for Spain: Torre la Higuera, Huelva, 22. iv
-09.V.1983, leg. J. Wolschrijn, genitalia slide R0464
(collection of senior author).
The form mundella is almost exclusively found along
the sandy coasts of the North Sea (the junior author
collected specimens up to 50 km inland in Jutland,
Denmark): S. Norway, Sweden (only one record from
the west coast), Denmark, Northwest Germany, Hol-
land, U.K., Ireland and France (Karsholt & Riedl loc.
cit.). Claims of mundella from other countries are
based upon misidentifications (see e.g. Karsholt 1995,
Karsholt & Huemer 1995). Records from Lithuania
(Ivinskis & Piskunov 1976) refer to B. similis, those
from Japan (Ueda et al. 1995) refer to B. svenssoni
(Park), or a closely related species.
Remarks
The respective lectotypes of B. umbrosella and B.
mundella are shown in figs. 9 and 10. Even though
their colours have bleached with time, the contrast
between the two moths is still remarkable and ex-
plains why they were considered separate species.
One has to realise, however, that the external features
of Bryotropha species can vary a great deal. The nom-
inate form of B. umbrosella with its dark, almost
black, forewing and contrasting white markings (figs.
1, 2), is typical for inland locations. In coastal areas
specimens often have a lighter colour owing to a more
or less heavy irroration with whitish scales (figs. 3, 4).
Variations also occur in B. mundella. Pure white spec-
imens (fig. 5) are rather rare. Most moths show a
weak (fig. 6) to heavy (fig. 7) irroration with darker,
greyish, scales, often disclosing the presence of costal
and tornai patches (see the lectotype in fig. 10). In
cases of a strong irroration with dark scales, the spec-
imens become greyish with prominent costal and tor-
nai patches (fig. 8). Compared with the B. umbrosella
in figs. 2 to 4, the dark B. mundella shown in fig. 8
can now be recognised for what it is: a very light form
of B. umbrosella.
The smooth transition of a nearly black B. umbrosel-
la into a nearly white B. mundella (figs. 1 -8) thus inval-
idates the last character separating B. mundella from B.
umbrosella. We thus have to conclude that B. mundella
is a coastal or ecological form of B. umbrosella.
The claim that moths of these two taxa differed in
wingshape was disproved by Richardson (1890). The
darker colour of the fringe in some B. mundella spec-
imens cause the wing itself to stand out strikingly and
appear shorter than it is. Taking the fringe into ac-
count, B. mundella has the same wingspan and wing-
shape as B. umbrosella.
Notes on synonymy
Gelechia umbrosella was described from one male
and three females collected in Poland, Zielona Gòra,
Glogów ('Glogau').
Gelechia mundella Douglas was described from an
unspecified number of specimens found in June on
sand hills in Great Britain, New Brighton, Cheshire.
The lectotype present in the bmnh was dissected a
long time ago ('Genitalia no. 139'), and the genital
slide can no longer be traced. However, a preparation
of a female from the same type-series (slide bmnh no.
25.320) confirmed that the genitalia are identical to
112
RuTTEN & Karsholt: Bryotropha umbrosella
Fig. 13. Lectotype of G. portlandicella; Great Britain; Port-
land, Dorset, Richardson coll. 1889 (dorc).
those of B. umbrosella.
Gelechia portlandicella was described from five spec-
imens collected in Gr. Britain, Portland, Dorset in
June 1888. Meyrick (1895) synonymised G port-
landicella with B. mundella. However, doubts on the
status of G. portlandicella remained (see e.g. Pierce &
Daltry 1938). We identified the original type series
preserved in the dorc as light forms of B. umbrosella.
Three specimens had been labeled 'co-types' and had
been dissected by N.H. Bennet in the British Museum
in 1937. The genitalia of the two males and one female
confirmed that the moths belong to B. umbrosella. We
have selected the female for the lectotype (fig. 13).
Bryotropha umbrosella fulvipalpella Joannis was de-
scribed from a long series of specimens ['en grand
nombre'] from France, Bretagne, Plouharnel. In these
moths, segment 2 of the labial palpus is orange. The
ZMUC possesses two specimens of B. umbrosellalabeWed
'Vannes, Bretagne'. They originate from Staudinger &
Bang-Haas and are probably part of Joannis' type se-
ries (although we can not prove this). Segment 2 of the
labial palpus of these moths indeed have an orange
haze. Though this feature is not found in specimens
from central or northern Europe, we consider this to
be within the variation of B. umbrosella. The status of
fidvipalpella as a form of B. umbrosella remains.
Gelechia anacampsoidella was described from an
unspecified number of specimens of both sexes from
Finland, Tvärminne. It was synonymed with B. um-
brosella by Hering (1926). Sattler (1960: 29) was of
the opinion that B. anacampsoidella should be regard-
ed as a subspecies of B. umbrosella. However, speci-
mens from Finland fall within the range of variation
of (dark) umbrosella specimens from Central and
northern Europe.
Benander (1961: 245) considered Tinea oppositella
Thunberg, 1794 a senior synonym of B. umbrosella.
However, oppositella of Thunberg is currently consid-
ered a misidentification of Alucita oppositella Fabri-
cius, 1775, a junior synonym of Borkhausenia minu-
tella (Linnaeus, 1758) (Oecophoridae).
Lempke (1976: 25) and Küchlein (1993: 272) list-
ed B. fuliginosella Snellen (1882) as a synonym of B.
umbrosella. A study of the type material of B. fuligi-
nosella preserved in rmnh proved it to belong to B.
similis (Karsholt & Kristensen 1995: 474).
Profound coastal or ecological variations are not
unique to B. umbrosella but are also found in B. affi-
nis. The latter species is dark coloured with yellowish
markings (fig. 1 1). In specimens from coastal areas the
forewing often is heavily irrorate with yellowish scales.
This sometimes gives rise to extreme light forms (fig.
12) with very distinct discal and plica! spots and near
to indistinct costal and tornai patches. Their yellowish
to brownish colour separates light forms of B. affinis
from light forms of B. umbrosella which are greyish or
whitish.
As pointed out by Karsholt & Skou (1987: 88), the
record of specimens of B. mundella in copula with B.
affinis (Larsen 1927: 97) is erroneous. All specimens
in question (at least as far as they are preserved in the
zmuc) belong to the nominate form of B. umbrosella.
Conclusions
During our study on B. mundella we noticed that
the genitalia of this moth are identical to those of B.
umbrosella and that the geographical distribution of
B. mundella is completely covered by that of B. um-
brosella. B. mundella is mainly found in coastal areas,
flying amongst nominate forms of B. umbrosella. Also
in their external features, we found all sorts of inter-
mediate forms linking an almost black B. umbrosella
to an almost white B. mundella. The evidence thus in-
dicates that B. mundella is a light, coastal or ecologi-
cal form of B. umbrosella.
Acknowledgements
We are indebted to the following persons for loan of
material for this study: R. de Peyer, Dorset County
Museum, Dorchester, U.K.; P. Falck, Holstebro,
Denmark; K. J. Huisman, Wezep, The Netherlands;
J. Kaila, Zoological Museum, University of Helsinki,
Finland; J. C. Koster, Callantsoog, The Netherlands;
J. Küchlein, Wageningen, The Netherlands; J. R.
Langmaid, Southsea, U.K.; Dr. E. J. van Nieukerken,
National Museum of Natural History, Naturalis, Lei-
den, The Netherlands; M. Shaffer, The Natural His-
tory Museum, London, U.K.; F. Vilhelmsen, Vanlose,
Denmark; J. B. Wolschrijn, Twello, The Netherlands.
Moreover we would like to thank L. Bot, Nieuw
Formerum, The Netherlands for instigating this
study. For the photographs we are indebted to M.
Shaffer and K. Tuck bmnh, and Sue Bunnewell, Nor-
113
Tijdschrift voor Entomologie, volume i4i, 1998
wich, U.K. We are grateful to P. Sterling (Wey-
mouth, UK) for help with obtaining the type series of
G. portlandicella, for comments on the manuscript
and for correcting the language.
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'Insecta Svedica', 1784-1794. - Opuscula Entomologica
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Douglas, J. W., 1849-52. On the British species of the genus
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Hering, M., 1924. Beitrag zur Kenntnis der Microlepido-
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Hering, M„ 1926. Ist Gelechis anacampsiodella Her. eine gute
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Karsholt, O., 1995. Kommentiertes Verzeichnis der Sym-
mocidae, Blastobasidae und Gelechiidae Ostdeutschlands
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Karsholt, O. & P. Huemer, 1995. Additions and corrections
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Karsholt, O. & N. P. Kristensen, 1995. [Review of] J. H.
Küchlein (collaborator: J. H. Donner): De Kleine Vlin-
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crolepidoptera. - Entomologica scandinavica 26: 473-474.
Karsholt, O. & T. Riedl, 1996. Gelechiidae, excl. Gnori-
moschemini. Pp. 103-113, 118-122, 310-312. - In: O.
Karsholt & J. Razowski (eds.), The Lepidoptera of Eu-
rope. - Stenstrup, 380 pp.
Karsholt, O. & P. Skou, 1987. Sommerfugle (Lepidoptera)
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Faunistiek van de Nederlandse Microlepidoptera. — Pu-
doc, Wageningen, 715 pp, 8 pis.
Larsen, C. S., 1927. Tillseg til Fortegnelse over Danmark. Mi-
crolepidoptera. - Entomologiske Meddeleser 17: 7-221.
Lempke, B. J., 1976. Naamlijst van de Nederlandse Lepi-
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Meyrick, E., 1895. A handbook of British Lepidoptera. -
London, vi + 843 pp.
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Pierce, F. N. & H. W. Daltry, 1938. Mniophaga: a new
genus of Gelechiidae, with reinstatement of portlandicel-
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(ed.), 1990, Keys to the Insects of the European Part of
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Received: 8 April 1998
Accepted: 15 June 1998
114
JOHN D. OSWALD
Department of Entomology, Texas A&M University, College Station, TX, U.S.A.
ANNOTATED CATALOGUE OF THE DILARIDAE
(INSECTA: NEURO PTERA) OF THE WORLD
Oswald, J. D., 1998. Annotated Catalogue of the Dilaridae (Insecta: Neuroptera) of the World.
-Tijdschrift voor Entomologie 141: 1 15-128. [issn 0040-7496]. Published 30 November 1998.
The neuropteran family Dilaridae is catalogued. Data on the status, primary type, type locality
and original place of publication are given for 67 valid species and 14 synonymous species-
group names. Status, type species, etymology and gender information are given for four valid
genera and seven synonymous genus-group names. Nepal is reported as a previously unrecog-
nized synonym of Dilar. Dilar grandis and Dilar marmoratus are new combinations from Rex-
avius. Berothella bannana is removed from the Dilaridae and transferred to the family Beroth-
idae as Berotba bannana, comb. n.
Dr. John D. Oswald, Department of Entomology, Texas A&M University, College Station,
TX 77843-2475 USA E-mail: j-oswald@tamu.edu.
Key Words. - Dilarinae, Nallachiinae, pleasing lacewings, systematics, taxonomy
The neuropteran family Dilaridae, 'pleasing lace-
wings', is a small family presently composed of 67
valid recent species, with a combined distribution en-
compassing parts of North and South America, Eu-
rope, Asia and Africa. Dilarids form a distinct clade
within the order Neuroptera, and are characterized by
males with pectinate antennae, females with elongate
ovipositors (a feature that they share with raphid-
iopterans and some mantispids), and distinctive de-
tails of the terminalia in both sexes. Dilarids are rela-
tively rare both in the field and in collections and are
of no known economic importance. Immatures are
associated with soil or dead wood. No fossil dilarids
are currently known. The purpose of the present
work is to provide a concise bibliographic, nomen-
clatural and taxonomie foundation for the family that
can serve the twin aims of providing an entry point
into the scattered dilarid literature and a stimulus for
further systematics study of the group.
History
The rarity of dilarids in the field is underscored by
the fact that the first known species, Dilar nevadensis,
does not appear in the scientific literature until 1838
(Rambur 1838, illustrated), and was not formally de-
scribed until four years later (Rambur 1 842) - a com-
paratively late date for a genus of insects that possess-
es several species with an aggregate distribution
stretching broadly across southern Europe. Dilar ne-
vadensis was treated at its inception as a new species
and genus, and it and its relatives have long been rec-
ognized as a distinctive group within the Neuroptera.
The genus Dilar was originally placed by Rambur in
his 'Tribu Semblides', together with several genera
now placed in the orders Megaloptera and Raphidio-
ptera. Soon thereafter, Newman (1853), in his reclas-
sification of the Neuroptera sensu lato, provisionally
placed the 'Dilaridae' as a division of his group Ste-
goptera, subgroup Corydalina, along with taxa now
attributed to the Ithonidae and Megaloptera. New-
man's classification was not widely adopted and his
name Dilaridae appears not to have come into gener-
al use. By the late 1800's Dilar was recognized as a
neuropteran (planipennian) genus, and was typically
included within a broad concept of the family He-
merobiidae. During the first two decades of the twen-
tieth century, the taxonomie concept of the Hemero-
biidae was radically altered. Many genera and groups
of genera were formally aggregated or reaggregated
into family-group taxa during this period, and several
of these were widely accorded full family rank.
Among these groups was the Dilaridae, which was
treated as a tribe (Dilarini, e.g., Navâs 1914), sub-
family (Dilarinae, e.g., Banks 1913) and full family
(Dilaridae, e.g., Handlirsch [1906]) during this time.
Current usage accords dilarids family rank, with two
subfamilies, the Dilarinae and Nallachiinae.
Navâs ([1909a], 1914), in the earliest revisionary
studies of the group, synthesized the prior descriptive
work of nineteenth century authors and his own early
twentieth century work on the fauna of Spain and
115
Tijdschrift voor Entomologie, volume hi, 1998
neighbouring regions. In his 1914 review of the fami-
ly for the Genera Insectorum, Navâs recognized seven
genera and 26 species. Of these, 21 species, but only 2
genera, are presently considered valid. During the
Navasian era heavy emphasis was placed on venation-
al traits as diagnostic characters for the identification
and justification of new neuropteran taxa at all ranks.
During this period, however, the full extent of vena-
tional variation was frequently not well known for
taxa established at both the specific and generic levels.
Subsequent réévaluation of generic limits, based prin-
cipally on male terminalic traits, has lead to a reduc-
tion in the number of accepted dilarid genera, and it
is now widely recognized that considerable venational
plasticity exists in many dilarid species. Characters of
the male terminalia have now come to replace vena-
tional traits as the characters of choice for delimiting
dilarid taxa, especially species. Although male termi-
nalic characters were figured as early as 1909 (Navâs
[1909a]), the added effort and difficulties of studying,
describing and figuring this complex character system
hindered its use and the full recognition of its value
during the early twentieth century. As in other neu-
ropterid groups, however, dilarid studies profited
from the growing realization throughout the 1930's
that the male terminalia provided a significant new
source of novel, stable and reliable species-level char-
acters. Subsequent reviews and revisions of dilarid
taxa (e.g., Carpenter 1940, 1947, Nakahara 1955b)
made extensive use of male terminalic traits, and
modern descriptions and revisions (e.g., Adams 1970,
Aspöck et al. 1980, Monserrat 1988b) rely heavily on
male terminalic characters to delimit species.
Future Research
The most pressing current need in dilarid system-
atics is for a comprehensive revision of Old World di-
larine species. The last complete revisions covering
the Old World are the seriously outdated works of
Navâs ([1909a], 1914). Since that time, the number
of nomenclaturally valid Old World dilarid species
has approximately tripled. The European dilarid fau-
na has recently been the subject of several good revi-
sionary treatments (Aspöck et al. 1980, Monserrat
1988b), but the lack of a synthetic work for the Ori-
ental region and the adjacent parts of central and east-
ern Asia renders positive identification of material
from these areas extremely difficult. The size of such
a project (ca. 50 species) would be quite appropriate
for a Ph.D. -level dissertation. Questions of interest
include:
(i) How variable are male terminalic traits among
Old World dilarids?,
(ii) What monophyletic subgroups can be recog-
nized within this fauna?, based on what characters?,
(iii) What is(are) the phylogenetic relationship (s)
between Old and New World dilarids?,
(iv) How old is the dilarid clade?,
(v) Why are there no known dilarid fossils?,
(vi) Why are dilarids absent from Australia?
Any study of the higher phylogeny of the family
should include representatives of New World Nal-
lachius, Old World Dilar and Berothella and the actu-
al or possible Old World nallachiine taxa Nallachius
krooni (southern Africa), Nallachius ponomarenkoi
(Vietnam) and Neonallachius annandalei (India).
Biology
Dilarid biology is poorly known. The larvae of only
five species are known, only four of which have been
described. The best known of these is the Nearctic-
Neotropical species Nallachius americanus, whose lar-
vae have been reported from under the bark of both
recently- and long-dead broadleaf trees, where they
undoubtedly feed on soft-bodied arthropod prey that
occur in the same microhabitat (Gurney 1947,
MacLeod & Spiegler 1961). Larvae of Dilar turcicus
and Dilar septentrionalis have been collected from soil
samples, Ghilarov (1962) and Makarkin (pers.
comm., larvae not described), respectively. The larvae
of two additional species, Nallachius krooni and Dilar
pumilus, have been reared from eggs laid by captive
females, see Minter (1992) and Monserrat (1988b).
Until recently, all known adult dilarids were fully
macropterous and presumed to be volant. However,
Pantaleoni & Letardi (1996) report the capture of a
brachypterous Dilar female that is probably attribut-
able to Dilar parthenopaeus.
Keys
A key is provided below to the two subfamilies of
the Dilaridae. The subfamily Nallachiinae currently
contains only a single valid genus, Nallachius. The
three nomenclaturally valid dilarine genera listed in
the following catalogue are widely considered only
doubtfully distinct. For this reason, a key to these taxa
has not been attempted. The construction of a reli-
able key to supraspecific taxa within the Dilarinae
cannot proceed until revisionary studies that identify
more rigorously diagnosable subgroups within this
taxon have been completed. Bibliographic references
to published species-level keys in the genera Dilar and
Nallachius are cited below under their respective cata-
logue entries.
116
Oswald: Catalogue of the Dilaridae
Key to the subfamilies of the Dilaridae
(after Adams 1970, New 1989)
1. Forewing MA branching from R basal to fork
Rl-Rs (i.e., as a branch from the 'radial stem'; or,
rarely, MA entirely free from R); male antenna
with more than 3 apical flagellomeres that lack
lateral processes [Europe, northern Africa, Asia]..
Dilarinae
+ Forewing MA branching from R distal to fork
Rl-Rs (i.e., as a branch from RS); male antenna
never with more than 3 apical flagellomeres that
lack lateral processes [North and South America,
southern Africa, Asia] Nallachiinae
Catalogue
Format
All known family-, genus- and species-group names
that have been applied to taxa presently placed within
the family Dilaridae are treated in the catalogue be-
low. Information on taxon diversity and distribution,
important systematics literature and biology and im-
mature stages is summarized for all valid family- and
genus-group taxa. Genus-group name records pro-
vide, additionally, data on type species, etymology
and gender, with appropriate bibliographic refer-
ences. Species citations provide information on origi-
nal publication, distribution, type locality and prima-
ry type kind, sex and depository (to the extent
known). Type locality data is presented in a standard-
ized format using current political subdivision names,
with verbatim quotes from the literature provided
parenthetically where confusion might arise. Latitude
and longitude coordinates are provided for all sites
that could be rigorously located. Coordinate data are
shown in brackets if they were derived from secondary
sources (e.g., maps or gazetteers), but are shown un-
bracketed if they were cited in the original description
of a species. Synonymical citations are given under the
subheading 'Synonymy'. Under the subheading 'Sta-
tus', a recent (where possible) authoritative work is
cited that uses the name in the nomenclatural form in
which it is treated in the catalogue. The third edition
of the International Code of Zoological Nomencla-
ture has been applied to questions of nomenclature,
and relevant articles and sections of the Code are ref-
erenced throughout the catalogue text.
Collection Acronyms
The following collection acronyms are used in the
catalogue to indicate type repositories:
ASPOCK Horst & Ulrike Aspöck, private collection,
Wien [=Vienna], Austria;
bau Beijing Agricultural University Insect
Collection, Beijing, China;
BMNH Natural History Museum [formerly the
British Museum (Natural History)],
London, England, United Kingdom;
iNBio Instituto de Biodiversidad, Santo Domingo
de Heredia, Costa Rica;
INPA Instituto Nacional de Pesquisas da
Amazonia, Manaus, Brazil;
ISNB Institut Royal des Sciences Naturelles de
Belgique, Brussels, Belgium;
IZASB Institute of Zoology, Academia Sinica,
Beijing, China;
MCNM Museo Nacional de Ciencias Naturales,
Madrid, Spain;
Mcz Museum of Comparative Zoology,
Cambridge, MA, USA;
MNHP Museum National d'Histoire Naturelle,
Paris, France;
monserrat Victor J. Monserrat, private collection,
Madrid, Spain;
MZB Museo Zoologia, Barcelona, Spain
MZUN Museo di Zoologia, Università di Napoli,
Napoli [=NapIes], Italy;
NCIP National Collection of Insects, Pretoria,
South Africa;
NHMB Naturhistorisches Museum Basel, Basel,
Switzerland;
NHMW Naturhistorisches Museum, Wien
[=Vienna] , Austria;
NSMT National Science Museum (Natural
History), Tokyo, Japan;
Nzsi Zoological Survey of India, National
Zoological Collection, Calcutta, West
Bengal, India;
PMY Peabody Museum of Natural History, Yale
University, New Haven, CT, USA;
real P. Real, private collection, Aix-en-Provence,
France;
ZMHA Zoologisches Museum für Hamburg,
Hamburg, Germany;
ZMHB Museum für Naturkunde, Humboldt
Universität, Berlin, Germany;
ZMUM Zoological Museum, Moscow State
University, Moscow, Russia;
ZSM Zoologische Staatssammlung, Munich,
Germany.
Family Dilaridae Newman, 1853
Dilaridae Newman, 1853 (as a 'division' of the Corydalina).
Type genus: Dilar Rambur, [1838].
Species and distribution. - 67 species; Eastern
United States south to Argentina, including the West
Indies (17 spp.), Oriental and southern Palearctic re-
gions (49 spp.), southern Africa (1 sp.). No dilarids
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Tijdschrift voor Entomologie, volume hi, 1998
are currently known from tropical Africa, Australia,
Oceania or extreme northerly or southerly latitudes.
No fossils attributable to the Dilaridae are currently
known.
Reviews, revisions and keys. - Navâs [1909a]
(World, revision, keys); Navâs 1914 (World, review,
keys); see also Dilar and Nallacbius below.
Biology and immature stages. - See Dilar and Nal-
lacbius below. The larva described and figured by
Takahashi (1942) as a dilarid is probably the larva of
a nevrorthid of the genus Nipponeurorthus, not a di-
larid. Tjeder (1937) compared the adult morphology
of dilarids and raphidiopterans.
Subfamily Dilarinae Newman, 1853
Dilarinae - Banks 1913:211 (as a subfamily of the Hemer-
obiidae). Type genus: Dilar Rambur, [1838].
Dilarini - Navis, 1914: 5 (as a tribe of the Dilaridae). Type
genus: Dilar Rambur, [1838].
Species and distribution. - 48 species; Oriental and
southern Palearctic regions.
Reviews, revisions and keys. - See Dilar below.
Biology and immature stages. - See Dilar below.
Genus Berothella Banks, 1934
Berothella Banks, 1934: 567. Type species: Berothella phan-
tomaBanks, 1934: 568, by monotypy. Etymology: Unex-
plained, probably Beroth- (< BerothfaJ, a berothid genus-
group name) — ella (< L. -ellus, a diminutive suffix), in
reference to the original supposed berothid affinities of
this genus. Gender: Feminine, from the gender appropri-
ate to the suffix -ella, Art. 30b. Status: Oswald & Penny
1991: 12.
Notes: Berothella was originally placed in the family
Berothidae; it was transferred to the family Dilaridae by
MacLeod and Adams ([1968]: 256).
Species and distribution. - 2 species; China and
continental Malaysia. A third species described in this
genus, Berothella bannana, belongs to the Berothidae,
see 'Taxa removed from the Dilaridae' below.
Reviews, revisions and keys. - None.
Biology and immature stages. -Unknown.
Berothella phantoma Banks, 1934 [Malaysia]
Berothella phantoma Banks, 1934: 568. Holorype, male,
BMNH. Type locality: Malaysia: Selangor: Bukit Kutu
(hill) [3°33'N 101°43'E]. Status: MacLeod & Adams
([1968]: 256).
Notes: Type citation by MacLeod & Adams ([1968]:
256). This species was originally described as a berothid.
It was confirmed as a dilarid by Kimmins in MacLeod &
Adams ([1968]: 256).
Berothella pretiosa Banks, 1939 [China]
Berothella pretiosa Banks, 1939: 469. Holorype, male, mcz.
Type locality: China: Kwangtung: Hainan (island), Ta-
han'. Status: MacLeod & Adams ([1968]: 256).
Notes: This species was originally described as a berothid. It
was confirmed as a dilarid by MacLeod & Adams ([1968]:
256).
Genus Dilar Rambur, [1838]
Dilar Rambur, [1838]: pi. 9. Type species: Dilar nevadensis
Rambur, [1838]: pi. 9, by monotypy. Etymology: From
Dilar, a river flowing west from the western edge of the
Sierra Nevada (mountains) of southern Spain, see Navâs
([1909a]: 628). Gender: Masculine, no originally attrib-
uted or implied gender, here considered masculine, Art.
30d. Status: Oswald & Penny 1991: 21. Known incorrect
subsequent spellings: Dillar, Dialar, Didar.
Cladocera Hagen, I860: 56, nomen nudum. Included spe-
cies: Cladocera marmorata Hagen, 1860: 56, nomen
nudum. Etymology: Unexplained, probably Clado- (<
Gr. klados, branch or twig) - -cera (< Gr. keras, horn), in
reference to the branched antennae, a characteristic of
male dilarids. Status: Oswald & Penny 1991: 21.
Notes: 'Cladocera marmorata Hoffm. Mus. Berol.' was
cited by Hagen (1860: 56) without description, defini-
tion or indication, and is therefore unavailable, Art. 12a.
The binomen Cladocera marmorata apparently originated
as a manuscript name that was recorded by Hagen from
the identification label of a specimen in the Hoffmansegg
Collection of the Museum ftir Naturkunde, Humboldt
Universität, Berlin. Hagen subsequently (1866b: 399)
synonymized Cladocera with Dilar., and C. marmorata
with Dilar nevadensis Rambur, without either name ever
having been made available.
Lidar Navâs, 1909b: 153. Type species: Dilar meridionalis
Hagen, 1866a: 295, by original designation. Etymology:
An anagram of Dilar, a dilarid genus-group name, see
Navâs (1909b: 153). Gender: Masculine, inferred from
the combination 'Lidar nemorosus , Art. 30d. Synonymy:
H. Aspöck et al. 1980: 187. Status: Oswald & Penny
1 99 1 : 21. Known incorrect subsequent spellings: Lider.
FuentenusNzvis, 1909b: 154. Type species: Dilar campestris
Navâs, 1903a: 380 {=Dilar saldubensis Navâs in Laguna,
1902), by original designation. Etymology: From the sur-
name of Reverend D. José Maria de la Fuente, Spanish
priest and entomologist, see Navâs (1909b: 155; [1909a]:
657). Gender: Masculine, no originally attributed or im-
plied gender, here considered masculine, Art. 30d. Syn-
onymy: H. Aspöck et al. 1980: 187. Status: Oswald &
Penny 1991:21.
Rexavius Navâs, [1909a]: 664. Type species: Dilar nietneri
Hagen, 1858b: 482, by subsequent designation by Navâs
1914: 10. Etymology: An anagram of Xaverius, from the
Latinized surname of Saint Francisco Javier, Spanish mis-
sionary to India and Japan, see Navâs ([1909a]: 664).
Gender: Masculine, inferred from the combination ' Rex-
avius jap onicus , Art. 30d. Synonymy: Kuwayama 1962:
376. Status: Oswald & Penny 1991: 21.
Notes: Rexavius was synonymized with Dilar without ex-
plicit comment by Kuwayama (1962: 376). Kuwayama's
synonymy was presumably based on his placement of the
species japonicus in the genus Dilar, rather than in Rexav-
ius, where it had been placed by Navâs (1909a: 665) as
one of the two species originally included in Rexavius.
The type species of Rexavius has also been transferred
back to Dilar from Rexavius by H. Aspöck & U. Aspöck
(1968: 3). Two additional species, grandis una marmora-
tus, also nominally exist in the genus Rexavius. Both are
here transferred to Dilar. I have examined syntypical ma-
118
Oswald: Catalogue of the Dilaridae
terial of both grandis and marmoratus from the mcz, and
both species clearly fall within the broad concept of Dilar
adopted in this catalogue.
NepalNavis, [1909a]: 661. syn. n. Type species: Nepal har-
mandi Navis, [1909a]: 661, by original designation. Ety-
mology: From Nepal, a region (later country) of the Indi-
an subcontinent that contains, or lies near, the type
localities of the two species originally included in this
genus, see Navâs ([1909a]: 661). Gender: Masculine, no
originally attributed or implied gender, here considered
masculine, Art. 30d.
Notes: All four species that have at one time or another
been placed in this genus have subsequently been re-
moved to Dilar, formosanus and kanoi by Nakahara
(1955b) and hornei and harmandihy H. Aspöck & U. As-
pöck (1968). Thus, although it has not previously been
listed as such, Nepal is currently a junior subjective syn-
onym of Dilar.
Species and distribution. - 45 species; Oriental and
southern Palearctic regions.
Reviews, revisions and keys. - Navâs 1903a (Spain,
review); Kuwayama 1921 (Japan, review, key); Naka-
hara 1955b (Japan & Taiwan, revision); Kuwayama
1962 (Japan, review, key); Aspöck et al. 1980 (Europe,
review, key); Minter 1986 (Africa, review); Dorokho-
va 1987 (European USSR, key); Monserrat 1988a
(Navâs species, revision); Monserrat 1988b (Iberian
Peninsula, revision, keys); Zakharenko 1988 (USSR,
review); Makarkin 1995 (Far Eastern Russia, key).
Biology and immature stages. - Ghilarov 1962
[turcicus: biology, larva, figs); Popov 1973 [turcicus:
egg, larva); Gepp 1984 [turcicus: larva, fig); Monser-
rat 1988b [pumilus: larva, figs); Gepp 1990 [turcicus:
egg, fig). The larva of Dilar septentrionalis is known
(Makarkin, pers. comm.), but undescribed.
Dilar algériens Navâs, 1909 [Algeria] Nomen
dubium
Dilar algériens Navas, [1909a]: 638. Holotype, male, mnhp.
Type locality: Algeria: Alger [36°50'N 3°00E, =Algiers,
=Argel (Spanish)] (as: 'Argel'). Status: Monserrat 1988a: 19.
Notes: See Legrand & Lachaise (1994: 87) for a detailed
account of the holotype; earlier type citations by Navas
(1925: 188) and Monserrat (1988b: 199).
Dilar aspersus C. Yang in Huang et al., 1988
[China]
Dilar aspersus C. Yang in Huang et al., 1988: 197. Holo-
type, male, bau. Type locality: China: Tibet: Nyingchi
County (as: 'Xizang: Nyingchi County'). Status: C. Yang
in Huang et al. 1988: 197.
Dilar bolivari Navte, 1903 [Algeria, Tunisia]
Dilar bolivari Navâs, 1903b: 116. Lectotype, male, mcnm,
designated by Monserrat (1988a: 16). Type locality: Al-
geria: Oran [35°45'N 0°38'W]. Status: Monserrat
1988a: 16.
Dilar caesarulus H. Aspöck & U. Aspöck, 1967
[Afghanistan]
Dilar caesarulus H. Aspöck &C U. Aspöck, 1967: 57. Holo-
type, male, aspock. Type locality: Afghanistan: 125 km
SW of Ghazni [G = 33°33'N 68°28'E]. Status: H. As-
pock & U. Aspöck 1967: 57.
Dilar corsicus Navâs, 1909 [France]
Dilar corsicus Navâs, [1909a]: 636. Holotype, female,
BMNH. Type locality: France: Corsica, 'La Foca' [=?Foce,
41°38'N 9°04'E]. Status: Monserrat 1988a: 17.
Notes: Type citation by Navâs (1925: 188).
Dilar budtzi Esben-Petersen, 1913: 27. Syntypes, male & fe-
male, repository unknown. Type locality: France: Corsica
(island). Synonymy: H. Aspöck et al. 1980: 188.
Dilar dissimilis Navâs, 1 903 [Spain]
Dilar dissimilis Navâs, 1903a: 374. Lectotype, maie, mnhp,
designated by Monserrat (1988b: 192). Type locality:
Spain: Zaragoza: Monasterio Santa Maria de Veruela
[4l°48'N 1°42"W], near Moncayo (mountain) (as:
'Veruela (Zaragoza) al pie del Moncayo'). Status: Mon-
serrat 1988b: 189.
Notes: See Legrand & Lachaise (1994: 88) for a detailed
account of the lectotype.
Dilar neposNavis, 1909b: 151. Lectotype, male, mnhp, des-
ignated by Legrand & Lachaise (1994: 90). Type locality:
Spain: Zaragoza: Muel [4l°28'N 1°04'W]. Synonymy:
H. Aspöck et al. 1980: 188. Status: Monserrat 1988: 189.
Notes: Earlier type citations by Navâs (1925: 188) and
Monserrat (1988b: 192).
Dilar kolbei Navâs, [1909a]: 635. Lectotype, male, mnhp,
designated by Navâs (1925: 188). Type locality: Spain:
Andaluda (region). Synonymy: H. Aspöck et al. 1980:
188. Status: Monserrat 1988: 189.
Notes: The type series of kolbei consisted of two male syn-
types, both of which were originally contained in the zmhb
(Navâs [1909a]: 635). One syntype, however, was appar-
endy retained by Navâs in his personal collection, and ulti-
mately came to be deposited in the mnhp. Navâs (1925:
188) clearly states that the specimen in the mnhp is the
'Type'. This statement constitutes a valid lectotype desig-
nation under Art. 74(b) of the Code, and fixes the mnhp
specimen as the lectotype of kolbei. The subsequent desig-
nation of the zmhb specimen as the lectotype by Legrand
and Lachaise (1994: 89) is therefore invalid, Art. 74a(i).
Navâs' ([1909a]: 635) statement: 'El tipo se halla en el mu-
seo de Berlin', is not a valid lectotype designation because it
does not identify a single specimen as the type — Navâs hav-
ing earlier clearly referred to two specimens in this collec-
tion ('He visto dos ejemplares que del museo de Berlin ...').
It might be argued that Navâs' 'tipo' statement referred to
the single specimen remaining in the zmhb after he re-
moved the second specimen of the type series to his own
personal collection. Under this interpretation, Navas' state-
ment would constitute a holotype designation (which
would then also invalidate the lectotype designation of Le-
grand & Lachaise). However, as Navâs gives no indication
that only one specimen remained in the zmhb at the time
of his writing, there is no concrete support for this interpre-
tation. See also the discussion by Monserrat (1988b: 192),
who treated the mnhp and zmhb specimens as syntypes.
Dilar distinctus Nakahara, 1955 [Taiwan]
Dilar distinctus Nakahara, 1955b: 139. Holotype, male,
NSMT. Type locality: Taiwan: Nantou: Sungkang (as:
Tattaka'). Status: Stange & Wang 1997: 49.
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Tijdschrift voor Entomologie, volume i4i, 1998
Dilar dochaner H. Aspöck & U. Aspöck, 1968
[Afghanistan]
Dilar dochanerW. Aspöck & U. Aspöck, 1968: 5. Holotype,
male, aspöck. Type locality: Afghanistan: Kabul: Khurd-
Kabul [34°24'N 69°24'E], SE of Kabul. Status: H. As-
pöck & U. Aspöck 1968: 5.
Dilar dongchuanus C. Yang, 1986 [China]
Dilar dongchuanus C. Yang, 1986: 155. Holotype, male,
bau. Type locality: China: Yunnan: Dongchuan
[26°24'N 103°08'E; =Tung-ch'uan, =Tungchwan,
=Hweitseh, =Tangdan]. Status: C. Yang 1986: 155.
Dilar duelli U. Aspöck & H. Aspöck, 1995 [France]
Dilar duelli U. Aspöck & H. Aspöck, 1995: 50. Holotype,
male, nhmw. Type locality: France: Var: Massif de l'Es-
térel, Col du Mistral, ENE of Fréjus, ca. 43°26'N 6°44'E.
Status: U. Aspöck & H. Aspöck 1995: 50.
Dilar formosanus (Okamoto & Kuwayama, 1 920)
[Taiwan]
Lidar formosanus Okamoto & Kuwayama, 1920: 341. Holo-
rype, female, repository unknown. Type locality: Taiwan:
'Arisan' [=?Chiayi Prefecture: Alishan or A-li-shan,
23°30'N 120°49'E]. Status: Stange & Wang 1997: 49.
Dilar geometroides H. Aspöck & U. Aspöck, 1968
[Nepal]
Dilar geometroides H. Aspöck & U. Aspöck, 1968: 3. Holo-
type, male, zsm. Type locality: Nepal: 'Prov. Nr. 3 East,
Sete'. Status: H. Aspöck & U. Aspöck 1968: 3.
Dilar grandis (Banks, 1931) [Malaysia (Sabah)]
comb. n.
Rexavius grandis Banks, 1931a: 413. Syntypes, male, mcz.
Type locality: Malaysia: Sabah: Borneo, Mt. Kinabalu,
Kamborangah.
Notes: See discussion under Rexavius above.
Dilar harmandi (Navâs, 1909) [India]
Nepal harmandi Navas, [1909a]: 661. Holotype, male,
MNHP. Type locality: India: West Bengal: Darjeeling
[27°02'N 88°20'E] (as: 'Darjeeling en el Himalaya). Sta-
tus: H. Aspöck & U. Aspöck 1968: 3.
Notes: See Legrand & Lachaise (1994: 88) for a detailed
account of the holotype; earlier type citation by H. As-
pöck & U. Aspöck (1968: 3).
Dilar hikosanus Nakahara, 1955 [Japan]
Dilar hikosanus Nakahara, 1955b: 137. Holotype, male,
probably in the nsmt. Type locality: Japan: Fukuoka /
Oita: Kyushu (island), Hiko-san (mountain) [33°29'N
130°58'E] (as: 'Hikosan, Kyushu' ). Status: Kuwayama
1962: 376.
Dilar hornei McLachlan, 1869 [India]
Dilar hornei McLachlan, 1869: 239. Syntype(s), male,
BMNH. Type locality: Northwestern India. Status: Ghosh
&Sen 1977:281.
Notes: Type citation by H. Aspöck & U. Aspöck (1968: 3).
Dilar indiens Monserrat, 1989 [India]
Dilar indicus Monserrat, 1989: 420. Holotype, male, nhmb.
Type locality: India: 'Cachemira, Rampur'. Status: Mon-
serrat 1989: 420.
Dilar japonicus McLachlan, 1883 [Japan]
Dilar japonicus McLachlan, 1883: 220. Holotype, male,
BMNH. Type locality: Japan: Fukushima: Honshu (is-
land), Fukushima [37°44'N 140°28'E] (as: 'Japan
(Fukushima in the main Island, ...'). Status: Kuwayama
1962: 376.
Dilar nohirae Nakahara, 1914: 297. Syntype(s), sex(es) un-
known, possibly in the nsmt. Type locality: Japan: Nara:
Yoshino [34°21'N 135°51'E] (as: Yoshino, Province Ya-
mato'). Synonymy: Nakahara 1955b: 134. Status: Ku-
wayama 1962: 376.
Dilar japonicus var. gracilis Kuwayama, 1921: 71.
Syntype(s), male, Okamoto collection (current repository
unknown). Type locality: Not fixed, see Notes. Syn-
onymy: Nakahara 1955b: 134. Status: Kuwayama 1962:
376.
Notes: Kuwayama (1921: 80) mentions three specimens
from southern Honshu (island), Japan, in the type series,
all in the Okamoto collection.
Dilar juniperi Monserrat, 1988 [Spain]
Dilar juniperi Monserrat, 1988b: 198. Holotype, male,
monserrat. Type locality: Spain: Jaén: Collado de los
Jardines [38°20'N 3°30'W]. Status: Monserrat 1988b:
198.
Dilar kanoi (Nakahara, 1955) [Taiwan]
Nepal kanoi Nakahara, 1955a: 6. Holotype, male, nsmt.
Type locality: Taiwan: Nantou: Sungkang (as: 'Tattaka').
Status: Stange & Wang 1997: 49.
Dilar kirgisus H. Aspöck & U. Aspöck, 1967
[Kirgizia]
Dilar kirgisus H. Aspöck & U. Aspöck, 1967: 59. Holotype,
male, aspock. Type locality: Kirgizia: Issyk-Kul (lake).
Status: Zakharenko 1988: 764.
Dilar lineolatus Navâs, 1909 [Turkey] Nomen
dubium
Dilar lineolatus Navâs, [1909a]: 645. Synrype(s), female,
ZMHB. Type locality: Turkey: Tekke [40°08'N 29°41'E] [as
'Tekke en la Turcomania'] . Status: Monserrat 1988a: 20.
Notes: Earlier type citation by H. Aspöck & U. Aspöck
(1968:3).
Dilar marmoratus (Banks, 1931) [Thailand] comb. n.
Rexavius marmoratus Banks, 1931b: 385- Syntypes, male &
female, mcz. Type locality: Thailand: Nakhon Si Tham-
marat [8°24'N 99°58'E], Khao Luang (mountain)
[8°31'N 99°47'E] (as: 'Peninsular Siam: Nakon Sri
Tamarat, Khao Luang').
Notes: See discussion under Rexavius above.
Dilar megalopterus C. Yang, 1986 [China]
Dilar megalopterus C. Yang, 1986: 154. Holotype, male,
bau. Type locality: China: Yunnan: Dongchuan
[26°24'N 103°08'E; =Tung-ch' uan, =Tungchwan,
=Hweitseh, =Tangdan]. Status: C. Yang 1986: 154.
Dilar meridionalis Hagen, 1866 [Andorra, France,
Spain]
Dilar meridionalis Hagen, 1866a: 295. Holotype, sex un-
known, repository unknown. Type locality: Spain: prob-
ably from the Sierra Nevada (mountains) near Granada.
120
Oswald: Catalogue of the Dilaridae
Status: Monserrat 1988b: 184.
Notes: The original description does not indicate where
the single specimen of the type series of meridionalis was
collected. Hagen (1866b: 402) subsequently cited the
species from Spain, and McLachlan (1869: 239) states
the type locality as 'the Sierra Nevada in the South of
Spain'.
Dilar pktus Navis, 1903a: 377. Holotype, female, mcnm.
Type locality: Spain: Madrid: near Madrid [M = 40°25'N
3°43'W]. Synonymy: Navâs 1909a: 652 (as a variety of
meridionalis). Status: Monserrat 1988b: 185.
Notes: Subsequently incorrectly cited as a new name by
Navâs ([1909a]: 652).
Lidar nemorosus Navâs, 1909b: 153. Syntype(s), male, repos-
itory unknown (see Notes). Type locality: Spain: Huesca:
Sierra de Guarà (mountains) near the Santuario de San
Cosme [SdSC = 42°15'N 0°16'W] (as: 'Sierra de Guarà
(Huesca), cerca del santuario de San Cosme'). Synonymy:
Navâs [1924]: 229. Status: Monserrat 1988b: 185.
Notes: Types presumed lost or destroyed. Probably for-
merly in the Navâs collection. Types not listed by Mon-
serrat (1985) as being in the remnants of the Navis col-
lection in the MZB.
Dilar mateui Real, 1968: 111. Holotype, male, real. Type
locality: France: Pyrénées-Orientales: 'Vallée de Nohèdes'
[Nohèdes (town) = ca. 1 1 km W of Prades, P=42°38'N
2°25'E]. Synonymy: H. Aspöck et al. 1980: 189. Status:
Monserrat 1988b: 185.
Dilar montanus C Yang, 1992 [China]
Dilar montanus C. Yang, 1992b: 441. Holotype, maie,
IZASB. Type locality: China: Szechwan [=Sichuan]: Zhon-
grewu, Xiangcheng [X = 28°54'N 99°40'E; = Hsiang-
ch'eng, = Hsiangcheng] . Status: C. Yang 1992b: 441.
Dilar nevadensis Rambur, 1838 [Spain]
Dilar nevadensis Rambur, [1838]: pi. 9. Syntype(s), sex(es)
unknown, isnb. Type locality: Spain: Granada: Sierra
Nevada (mountains), near Granada [G = 37°10'N
3°35'W] (as: 'aux environs de Grenade, dans les petits
bois des parties élevée de la Sierra-Nevada'). Status: Mon-
serrat 1988b: 182.
Notes: Type locality from Rambur (1842: 446). At least
two syn types are stated by Navis ([1909a]: 632) to be in
the isnb. Navis ([1909a]: 629) cites his earlier use of the
name ' nivatensis as an error for Dilar nevadensis Rambur.
Cladocera marmoratus Hagen, 1860: 56, nomen nudum.
Status: Hagen 1866b: 399.
Notes: 'Cladocera marmorata Hoffrn. Mus. Berol.' was
cited by Hagen (1860: 56) without description, defini-
tion or indication, and is therefore unavailable, Art. 1 2a.
The binomen Cladocera marmorata apparently originated
as a manuscript name that was recorded by Hagen from
the identification label of a specimen in the Hoffmansegg
Collection of the Museum für Naturkunde, Humboldt
Universität, Berlin. Hagen (1866b: 399) subsequently
synonymized Cladocera with Dilar, and C. marmorata
with Dilar nevadensis Rambur, without either name ever
having been made available.
Dilar nietneri Hagen, 1858 [Sri Lanka]
Dilar nietneri Hagen, 1858b: 482. Syntype(s), sex(es) un-
known, ZMHB. Type locality: Sri Lanka: 'Rainbodde'.
Status: H. Aspöck & U. Aspöck 1968: 3.
Notes: Type citations by Navâs ([1909a]: 664) and H.
Aspöck & U. Aspöck (1968: 3).
Dilar pallidusNakahzra, 1955 [Taiwan]
Dilar pallidus Nakahara, 1955b: 140. Holotype, male,
NSMT. Type locality: Taiwan: Nantou: Sungkang (as:
'Tattaka'). Status: Stange & Wang 1997: 49.
Dilar parthenopaeus A. Costa, 1855 [Italy]
Dilar parthenopaeus A. Costa, 1855: 19. Lectotype, male,
MZUN. Type locality: Italy: Campania: Salerno: Cava de'
Tirreni [40°42'N 14°42'E]. Status: H. Aspöck et al.
1980: 190.
Notes: Type locality information and lectotype designa-
tion from Pantaleoni (pers. comm.; data from manuscript
in preparation on Costa types).
Dilar pumilus Nzvis, 1903 [Spain]
Dilar pumilus Navâs, 1903a: 380. Holotype, male, mnhp.
Type locality: Spain: Murcia: near Cartagena [C =
37°36'N 0°59'W]. Status: Monserrat 1988b: 195.
Notes: See Legrand & Lachaise (1994: 90) for a detailed
account of the holotype; earlier type citations by Navis
(1925: 188) and Monserrat (1988b: 195).
Dilar pusillus C. Yang in Huang et al., 1992
[China]
Dilar pumilus C. Yang in Huang et al., 1988: 197. Holo-
type, male, probably in the bau or izasb. Type locality:
China: Tibet [=Xizang]: Medoge County, Beibeng. Sta-
tus: C. Yang 1992a: 379.
Notes: A junior primary homonym of Dilar pumilus
Navâs, 1903. The objective replacement name is Dilar
pusillus C. Yang, 1992.
Dilar pusillus C. Yang, 1992a: 379. Status: C. Yang 1992a:
379.
Notes: An objective replacement name for Dilar pumilus
C. Yang in Huang et al., 1988, nee Navâs, 1903.
Dilar saldubensis Navâs in Laguna, 1902 [Portugal,
Spain]
Dilar saldubensis Navâs in Laguna, 1902: 134. Neotype,
male, mnhp, designated by Monserrat 1988b: 188. Type
locality: Spain: Zaragoza: Zaragoza [4l°39'N 0°54'W].
Status: Monserrat 1988b: 187.
Notes: The original type series was formerly in the Navâs
collection. The specimens of this series are not present in
the remnants of the Navâs collection in the mzb (Mon-
serrat 1985), and are now presumed lost or destroyed. See
also Legrand & Lachaise (1994: 90) for a detailed account
of the neotype.
Dilar campestris Nzvis, 1903a: 380. Lectotype, male, mnhp,
designated by Legrand & Lachaise (1994: 88). Type local-
ity: Spain: Ciudad Real: Pozuelo de Calatrava [38°55'N
3°50'W]. Synonymy: H. Aspöck et al. 1980: 189. Status:
Monserrat 1988b: 187.
Notes: Earlier type citations by Navâs (1925: 188) and
Monserrat (1988b: 188).
Fuentenus lusitanicus Navâs, [1909a]: 660. Holotype, sex
unknown, zmhb. Type locality: Northern Portugal. Syn-
onymy: Navâs [1924]: 232 (as a synonym of campestris).
Status: Monserrat 1988b: 188.
Notes: Type in poor condition and missing its abdomen.
This species was considered to be a nomen dubium by
121
Tijdschrift voor Entomologie, volume mi, 1998
Monserrat (1988b: 188), but he accepted Navâs' ([1924]:
232) synonymy of lusitaniens with campestris.
Dilar septentrionalis Navâs, 1912 [China, Korea,
Russia]
Dilar septentrionalis Navâs, 1912: 420. Lectotype, male,
MNHP, designated by Monserrat (1988a: 20). Type local-
ity: Russia: Vladivostok [43°09'N 131°53'E]. Status:
Makarkin 1990: 38.
Notes: See Legrand & Lachaise (1994: 91) for a detailed
account of the lectotype.
Dilar similis Monserrat, 1989 [Pakistan]
Dilar similis Monserrat, 1989: 419. Holorype, male, nhmb.
Type locality: Pakistan: North-West Frontier: 'Salf-ui-
MalukSar'. Status: Monserrat 1989: 419.
Dilar sinicus Nakahara, 1957 [China]
Dilar sinicus Nakahara, 1957: 31. Holorype, male, probably
in the NSMT. Type locality: China: Shansi: between
Henglingkuan [35°25'N 111°36'E; =Heng-ling-kuan,
=Henglingguan] and Wangmaochen. Status: Nakahara
1957:31.
Dilar subdolus Navâs, 1932 [China] Nomen
dubium
Dilar subdolus Navâs, 1932: 921. Holorype, male, reposito-
ry unknown (see Notes). Type locality: China: Kiangsu:
Chinkiang [32°03'N 119°26'E; =Chen-chiang, =Chen-
kiang, =Zhenjiang]. Status: Monserrat 1988a: 22.
Notes: Holorype formerly in the Navâs collection, now
presumed lost or destroyed. Type not listed by Monserrat
(1985) as being in the remnants of the Navâs collection in
the MZB.
Dilar syriacusNzvis, 1909 [Syria] Nomen dubium
Dilar syriacus Navâs, [1909a]: 644. Holorype, male, mnhp.
Type locality: Lebanon: near Beirut [B = 33°52'N
35°30'E] (as: 'Siria, Creo que es de los alrededores de
Beirut'). Status: Monserrat 1988a: 19.
Notes: See Legrand & Lachaise (1994: 91) for a detailed
account of the holorype.
Dilar taiwanensis Banks, 1937 [Taiwan]
Dilar taiwanensis Banks, 1937: 276. Syntype(s), sex(es) un-
known, probably in the mcz. Type locality: Taiwan: Chi-
ayi [=Chia-i]: Alishan [=A-li-shan, 23°30'N 120°49'E]
(as: 'Arizan'). Status: Stange & Wang 1997: 49.
Dilar tibetanus C. Yang, 1987 [China]
Dilar tibetanus C. Yang, 1987: 197. Holotype, male, proba-
bly in the bau or izasb. Type locality: China: Tibet:
Borni Co., Yigang. Status: C. Yang 1987: 197.
Dilar turcicus Hagen, 1858 [Southern Europe from
Yugoslavia east to Dagestan (Russia)]
Dilar turcicus Hagen, 1858a: 129. Holotype, sex unknown,
repository unknown. Type locality: Turkey. Status: H.
Aspöck et al. 1980: 190.
Dilar corcyraeus Navâs, [1909a]: 642. Holorype, female,
NHMW. Type locality: Greece: Corfu (island). Synonymy:
H. Aspöck et al. 1980: 190. Status: Monserrat 1988a: 19.
Dilar vartianorum H. Aspöck & U. Aspöck, 1967
[Afghanistan]
Dilar vartianorum H. Aspöck & U. Aspöck, 1967: 57.
Holotype, male, aspock. Type locality: Afghanistan:
Nuristan, 25 km N of Barikot [B = 35°18'N 71°36'E].
Status: H. Aspöck & U. Aspöck 1967: 57.
Dilar vietnamensis Zakharenko, 1991 [Vietnam]
Dilar vietnamensis Zakharenko, 1991: 142. Holotype, male,
ZMUM. Type locality: Vietnam: 'Onang Ninh, Dongkho'.
Status: Zakharenko 1991: 142.
Dilar wangi C. Yang, 1 992 [China]
Dilar wangi C. Yang, 1992b: 441. Holotype, male, izasb.
Type locality: China: Yunnan: Wengshui, Zhongdian [Z
= 27°46'N 99°45'E; =Chung tien]. Status: C. Yang
1992b: 441.
Dilar yunnanus C. Yang, 1986 [China]
Dilar yunnanus C. Yang, 1986: 154. Holotype, male, bau.
Type locality: China: Yunnan: Dehong Dai-Jingpo ^Te-
ilung Shan-kachin] Autonomous District: Ruili
[24°01'N 97°52'E; =Jui-li]. Status: C. Yang 1986: 154.
Genus Neonallachius Nakahara, 1 963
Neonallacbius Nakahara, 1963: 77. Type species: Neonal-
lachius annandalei Nakahara, 1963: 77, by original desig-
nation. Etymology: Unexplained, probably Neo- (< Gr.
neos, new) — nallachius (< Nallachius, a dilarid genus-
group name). Gender: Masculine, no originally attrib-
uted or implied gender, here considered masculine, Art.
30d. Status: Oswald & Penny 1991: 42.
Species and distribution. - 1 species; India, Sri
Lanka; see Hynd (1992), Nakahara (1963).
Reviews, revisions and keys. - None.
Biology and immature stages. - Unknown.
Neonallachius annandalei Nakahara, 1 963 [India]
Neonallachius annandalei Nakahara, 1963: 77. Holotype,
male, nzsi. Type locality: India: Orissa: Barkuda Island,
Chilka Lake, near Ganjam [G = 19°28'N 85°05'E]. Sta-
tus: Adams 1970:7.
Subfamily Nallachiinae Navâs, 1914
Nallachini Navâs, 1914: 11 (as a tribe of the Dilaridae).
Type genus: NallachiusNuvis, 1909. Notes: Name incor-
rectly formed from the type genus.
Nallachiini - Carpenter, 1947: 100 (as a tribe of the Dilari-
dae). Type genus: Nallachius Navâs, 1909.
Nallachiinae - Adams, 1970: 8 (as a subfamily of the Dilar-
idae). Type genus: Nallachius Navâs, 1909.
Species and distribution. - See Nallachius below.
Reviews, revisions and keys. — See Nallachius be-
low.
Biology and immature stages. - See Nallachius be-
low.
Genus Nallachius Navâs, 1909
Nallachius Navâs, [1909a]: 666. Type species: Dilar prestoni
McLachlan, 1880: 39, by subsequent designation by
122
Oswald: Catalogue of the Dilaridae
Navâs 1914: 11. Etymology: An anagram of Lachlanius,
from the Latinized surname of Robert McLachlan, Eng-
lish entomologist, see Navâs ([1909a]: 665). Gender:
Masculine, inferred from the combination ' Nallachius
americanus', Art. 30d. Status: Oswald & Penny 1991: 39.
Nulema Navâs, 1914: 12. Type species: Nulema championi
Navâs, 1914: 12, by monotypy. Etymology: An arbitrary
combination of letters, see Navâs (1914: 12). Gender:
Feminine, no originally attributed or implied gender,
here considered a natural Latin feminine, Art. 30d. Syn-
onymy: Adams 1970: 8. Status: Oswald & Penny 1991:
39. Known incorrect subsequent spellings: Nurema.
Neodilar Carpenter, 1947: 107. Type species: Dilar hermosa
Banks, 1913: 220, by original designation. Etymology:
Unexplained, probably Neo- (< Gr. neos, new) - -dilar (<
Dilar, a dilarid genus-group name). Gender: Masculine,
no originally attributed or implied gender, here consid-
ered masculine, Art. 30d. Synonymy: Adams 1970: 8.
Status: Oswald & Penny 1991: 39.
Species and distribution. - 19 species; Eastern Unit-
ed States south to Argentina, including the West Indies
(17 spp.), southern Africa (1 sp.) and Vietnam (1 sp.);
see Adams (1970), Hoffman (1990), Maes & Flint
(1994), Penny ([1978], [1982]), Penny et al. (1997).
Reviews, revisions and keys. - Carpenter 1940
(Nearctic, review); Carpenter 1947 (New World, re-
view); Adams 1970 (New World, revision, key); Pen-
ny [1982] (Amazon Basin, review, key).
Biology and immature stages. - Steyskal 1944
{americanus: biology); Gurney 1947 {americanus: bi-
ology, egg, larva, pupa, figs); Peterson 1967 {ameri-
canus: larva, fig); MacLeod & Spiegler 1961 {ameri-
canus: biology, egg, larva); Tauber 1991 {americanus:
larva, fig); MacLeod 1964 {americanus: larva, figs);
Gepp 1984 {americanus: larva, fig); Minter 1992
{krooni: egg, larva, figs).
Nallachius adamsi Penny, 1982 [Brazil]
Nallachius adamsi Penny, [1982]: 385. Holotype, male,
INPA. Type locality: Brazil: Amazonas: Manaus [3°06'S
60°00'W]. Status: Penny [1982]: 385.
Nallachius americanus (McLachlan, 1881) [Eastern
USA south to Venezuela, West Indies]
Dilar americanus McLachlan, 1881: 55. Holotype, female,
mcz. Type locality: USA: Kentucky: Edmonson Co.: Bee
Spring [37°17'N 86°17'W]. Status: Penny et al. 1997: 63.
Notes: Type citation by Adams (1970: 27).
Nallachius bruchi Navâs, 1 923 [Argentina]
Nallachius bruchi Navâs, 1923: 195. Holotype, female,
repository unknown (see Notes). Type locality: Argenti-
na: Cordoba: Alta Gracia [31°42'S 64°25'W]. Status:
Penny [1978]: 30.
Notes: Holotype formerly in the Navâs collection, now
presumed lost or destroyed. Type not listed by Monserrat
(1985) as being in the remnants of the Navâs collection in
the MZB.
Nallachius championi (Navâs, 1914) [Guatemala]
Nulema championi Navâs, 1914: 12. Lectotype, male,
BMNH, designated by Adams (1970: 22). Type locality:
Guatemala: Cerro Zunil [14°44'N 91°27"W]. Status:
Penny [1978]: 30.
Nallachius dicolor Adams, 1970 [Argentina, Brazil]
Nallachius dicolor Adams, 1970: 19. Holotype, male, pmy.
Type locality: Brazil: Santa Catarina: Nova Teutonia,
27°H'S 52°23'W. Status: Penny [1978]: 30.
Nallachius hermosa (Banks, 1913) [Colombia]
Dilar hermosa Banks, 1913: 220. Holotype, female, mcz.
Type locality: Colombia: Cundinamarca: Cordillera Ori-
ental, Pacho [5°09'N 74°08'W]. Status: Penny [1978]:
30. Notes: Type citation by Adams (1970: 25).
Nallachius infiiscatus Penny, 1982 [Brazil]
Nallachius infiiscatus Penny, [1982]: 386. Holotype, male,
INPA. Type locality: Brazil: Amazonas: Reserva Ducke
[2°54'S 59°57'W], 26 km N of Manaus [3°07'S
60°02'W]. Status: Penny [1982]: 386.
Nallachius krooni Minter, 1 986 [Malawi, Namibia,
South Africa]
Nallachius krooni Mimet, 1986: 88. Holotype, male, nop.
Type locality: South Africa: Transvaal: Wylliespoort,
22°55'S 29°56'E. Status: Minter 1986: 88.
Nallachius limai Adams, 1970 [Brazil]
Nallachius limai Adams, 1970: 23. Holotype, male, pmy.
Type locality: Brazil: Santa Catarina: Nova Teutonia,
27°1 l'S 52°23'W. Status: Penny [1978]: 30.
Nallachius loxanus Navâs, 1911 [Ecuador]
Nallachius loxanus Navâs, 1911: 219. Holotype, male,
MNHP. Type locality: Ecuador: Loja: Loja [3°59'S
79°16W] (as: 'Equateur, Loja'). Status: Penny [1978]: 30.
Notes: See Legrand & Lachaise (1994: 89) for a detailed
account of the holotype.
Nallachius maculatus Penny, 1982 [Brazil]
Nallachius maculatus Penny, [1982]: 389. Holotype, male,
INPA. Type locality: Brazil: Rondonia: 48 km E of Porto
Velho [PV = 8°45'S 63°54"W]. Status: Penny [1982]:
389.
Nallachius ovalis Adams, 1970 [Brazil]
Nallachius ovalis Adams, 1970: 17. Holotype, male, pmy.
Type locality: Brazil: Santa Catarina: Nova Teutonia,
27° 1 l'S 52°23'W. Status: Penny [1978]: 30.
Nallachius par keriVenny, 1994 [Costa Rica]
Nallachius parkeri Penny, 1994: 309. Holotype, male, in-
bio. Type locality: Costa Rica: Guanacaste: 3 km SE of
Rio Naranjo (town) [RN = 10°41'N 85°06'W]. Status:
Penny 1994: 309.
Nallachius phantomellus Adams, 1970 [Brazil]
Nallachius phantomellus Adams, 1970: 12. Holotype, male,
pmy. Type locality: Brazil: Mato Grosso do Sul: Rio
Caragualà, 21°48'S, 52°27'W. Status: Penny [1978]: 30.
Nallachius ponomarenkoi Zakharenko, 1 99 1
[Vietnam]
Nallachius ponomarenkoi Zakharenko, 1991: 143. Holo-
type, male, zmum. Type locality: Vietnam: Kien Giang:
'Tho Tu'. Status: Zakharenko 1991: 143.
Nallachius prestoni (McLachlan, 1880) [Brazil]
Dilar prestoni McLachlan, 1880: 39. Holotype, male, bmnh.
123
Tijdschrift voor Entomologie, volume i4i, 1998
Type locality: Brazil: Rio de Janeiro: near Rio de Janeiro
[RdJ = 22°53'S 43°17'W]. Status: Penny [1978]: 30.
Notes: Type citation by Adams (1970: 17).
Nallachius pulchellus (Banks, 1938) [Southwestern
USA, Cuba, Costa Rica]
Dilar {Nallachius) pulchellus Banks, 1938: 289. Holotype,
maie, mcz. Type locality: Cuba: Cienruegos: Soledad,
near Cienruegos [C = 22°10'N 80°27"W]. Status: Penny
étal. 1997:62.
Notes: Type citation by Adams (1970: 14).
Nallachius pupillus (Navâs, 1930) [Paraguay]
Nulema pupillusNavâs, 1930: 62. Holotype, male, zmha (see
Notes). Type locality: Paraguay: Cordillera: San Bernardi-
no [25°16'S° 57T6W]. Status: Penny [1978]: 30.
Notes: Type citation by Adams (1970: 29). Type pre-
sumed destroyed in the Hambutg Museum during
WWII.
Nallachius reductus Carpenter, 1 947 [Paraguay]
Nallachius reductus Carpenter, 1947: 104. Holotype, male,
mcz. Type locality: Paraguay: Ilalyria (as: 'Ualyaia,
Paraguay'). Status: Penny [1978]: 30.
Notes: Type citation by Adams (1970: 15).
Taxa removed from the Dilaridae
Berotha bannana (C. Yang, 1986) [China] comb. n.
Berothella bannana C. Yang, 1986: 156. Holotype, female,
bau. Type locality: China: Yunnan: Xishuangbanna Dai
[=Hsi-shuang-pan-na Thai] Autonomous District,
Menghai [21°59'N 100°35'E; = Meng-hai].
Notes: Yang's figure clearly shows that this species be-
longs in the Berothidae, where it appears to be a species of
the genus Berotha (U. Aspöck, pers. comm.). The species
is here transferred to the Berothidae under the new com-
bination Berotha bannana.
Acknowledgments
I thank Ulrike Aspöck, Vladimir Makarkin, Victor
Monserrat and Roberto Pantaleoni for answering my
queries to a variety of questions, as attributed in the
text above. Horst Aspöck, Ulrike Aspöck, Vladimir
Makarkin, Victor Monserrat and Norm Penny kind-
ly reviewed an earlier draft of the manuscript.
Indices
(valid names italicized)
Genus-group name valid genus
Berothella Banks Berothella
Cladocera Hagen Dilar
Dilar Rambur Dilar
Fuentenus Navâs Dilar
Lidar Navâs Dilar
Nallachius Navâs Nallachius
Neodilar Carpenter Nallachius
Neonallachius Nakahara. Neonallachius
Nepal Navâs Dilar
Nulema Navâs Nallachius
Rexavius Navâs Dilar
Species-group name valid species
adamsi Penny Nallachius adamsi
algériens Navâs Dilar algériens
amerìcanus McLachlan .... Nallachius americanus
annandalei Nakahara . . . Neonallachius annandalei
aspersus C. Yang in Huang et al. ... Dilar aspersus
bannana C. Yang Berothella bannana
bolivari Navâs Dilar bolivari
bruchi Navâs Nallachius bruchi
budtzi Esben-Petersen Dilar corsicus
caesarulus H. Aspöck & U. Aspöck Dilar caesarulus
campestris Navâs Dilar saldubensis
championi Navâs Nallachius championi
corcyraeus Navâs Dilar turcicus
corsicus Navâs Dilar corsicus
dicolor Adams Nallachius dicolor
dissimilis Navâs Dilar dissimilis
distinctus Nakahara Dilar distinctus
dochaner H. Aspöck & U. Aspöck . Dilar dochaner
dongchuanus C. Yang Dilar dongchuanus
duelli U. Aspöck & H. Aspöck Dilar duelli
formosanus Okamoto & Kuwayama
Dilar formosanus
geometroides H. Aspöck & U. Aspöck
Dilar geometroides
gracilis Kuwayama Dilar japonicus
grandis Banks Dilar grandis
harmandi Navâs Dilar harmandi
hermosa Banks Nallachius hermosa
hikosanusNakahava. Dilar hikosanus
hornei McLachlan Dilar hornei
indicus Monserrat Dilar indicus
infuscatus Penny Nallachius infuscatus
japonicus McLachlan Dilar japonicus
juniperi Monserrat Dilar juniperi
kanoi Nakahara Dilar kanoi
kirgisus H. Aspöck & U. Aspöck . . . Dilar kirgisus
kolbei Navâs Dilar dissimilis
krooni Minter Nallachius krooni
limai Adams Nallachius limai
lineolatus Navâs Dilar lineolatus
loxanus Navâs Nallachius loxanus
lusitanicus Navâs Dilar saldubensis
macidatus Penny Nallachius maculatus
marmoratus Banks Dilar marmoratus
marmoratus Hagen Dilar nevadensis
mateui Real Dilar meridionalis
megalopterus C. Yang Dilar megalopterus
meridionalis Hagen Dilar meridionalis
montanus C. Yang Dilar montanus
nemorosus Navâs Dilar meridionalis
124
Oswald: Catalogue of the Dilaridae
nepos Navâs Dilar dissimilis
nevadensis Rambur Dilar nevadensis
niemeri Hagen Dilar nietneri
nohirae Nakahara Dilar japonicus
ovalis Adams Nallachius ovalis
pallidus Nakahara Dilar pallidus
parkeri Venny Nallachius parkeri
parthenopaeus A. Costa Dilar parthenopaeus
phantoma Banks Berothella phantoma
phantomellus Adams Nallachius phantomellus
pictus Navâs Dilar meridionalis
ponomarenkoi Zakharenko Nallachius ponomarenkoi
prestoni McLachlan Nallachius prestoni
pretiosa Banks Berothella pretiosa
pulchellus Banks Nallachius pulchellus
pumilus C. Yang in Huang et al. ... Dilar pusillus
pumi lus Navâs Dilar pumilus
pupillus Navâs Nallachius pupillus
pusillus (2. Yang Dilar pusillus
reductus Carpenter Nallachius reductus
saldubensis Navâs in Laguna .... Dilar saldubensis
septentrionalis Navâs Dilar septentrionalis
similis Monserrat Dilar similis
sinicus Nakahara Dilar sinicus
subdolus Navâs Dilar subdolus
syriacus Navâs Dilar syriacus
taiwanensis Banks Dilar taiwanensis
tibetanus C. Yang Dilar tibetanus
turcicus Hagen Dilar turcicus
vartianorum H. Aspöck & U. Aspöck
Dilar vartianorum
vietnamensis Zakharenko Dilar vietnamensis
wangi C. Yang Dilar wangi
yunnanus C. Yang Dilar yunnanus
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128
Tijdschrift voor Entomologie
Volume 141 , no. I
Articles
I D.J. Bickel
Australian, Melanesian, and Micronesian Acropsilus Mik (Diptera: Dolichopodidae).
19 S.-W. Choi
Systematics of the genus Heterothera Inoue (Lepidoptera, Geometridae: Larentiinae)
49 H. R. Feijen
Teleopsis Rondani (Diptera, Diopsidae): generic review and the ferrugìnea group
from Sri Lanka.
65 S. Ingrisch
A review of the Elimaeini of Western Indonesia, Malay Peninsula and Thailand (Ensi-
fera, Tettigoniidae, Phaneropterinae).
109 T. Rutten & O. Karsholt
Bryotropha mundella (Douglas): a new synonym of Bryotropha umbrosella (Zeiler)
(Lepidoptera Gelechiidae).
115 J.D.Oswald
Annotated catalogue of the Dilaridae (Insecta: Neuroptera) of the World.
Book reviews
18 Zlata S. Gershenson & Sandrine A. Ulenberg, 1998. The Yponomeutinae (Lepido-
ptera) of the World exclusive of the Americas. • Menno Schilthuizen & Henk Val-
lenduuk, 1998. Kevers op kadavers. • Johan van Zoest (ed.), 1998. Biodiversiteit.
[E.J. van Nieukerken]
48,64 Orthoptera sounds: D.R. Ragge & W.J. Reynolds, 1998. The songs of the Grass-
hoppers and Crickets of Western Europe. • D.R. Ragge & W.J. Reynolds, 1998. A
sound guide to the Grasshoppers and Crickets of Western Europe. • H. Bellmann,
1993a. Heuschrecken beobachten, bestimmen. • H. Bellmann, 1993b. Die Stimmen
der heimischen Heuschrecken. • H. Bellmann & G. Luquet, 1995. Guide des saute-
relles, grillons et criquets d'Europe occidentale. • F.-R. Bonnet, 1995. Guide sonore
des sauterelles, grillons et criquets d'Europe occidentale. • R.M.J.C Kleukers, E.J.
van Nieukerken, B. Odé, L.P.M. Willemse & W.K.R.E. van Wingerden, 1997. De
sprinkhanen en krekels van Nederland (Orthoptera). • B. Odé, 1997. De zingende
sprinkhanen en krekels van de Benelux. [E. J. van Nieukerken]
© Nederlandse Entomologische Vereniging, Amsterdam
Published 30 November 1998 ISSN 0040-7496
Volume 141, no. 2, 1998
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Ontwerpers B.V., Aad Derwort, 's-Gravenhage
Robert S. ANDERSON
Entomology-Canadian Museum of Nature, Ottawa
NEW SPECIES OF SICODERUS VANIN FROM THE
VIRGIN ISLANDS (COLEOPTERA: CURCULIONIDAE;
CURCULIONINAE; OTIDOCEPHALINI)
Anderson, R. S. 1999. New species of Sicoderus Vanin from the Virgin Islands (Coleoptera:
Curculionidae; Curculioninae; Otidocephalini). — Tijdschrift voor Entomologie 141 [1998]:
129-135, figs. 1-1 2. [issN 0040-7496]. Published 1 March 1999.
Three new species of the genus Sicoderus Vanin from the Virgins Islands of the Greater Antilles
are described. The three new species are Sicoderus ivieorum sp. n., S. hirsutiventris sp. n., and 5.
vanini sp. n. They are placed in the S. tinamus species group and compared with known West
Indian Sicoderus. A key to all Sicoderus in the West Indies (including Florida) is presented.
Robert S. Anderson, Entomology-Canadian Museum of Nature, P.O. Box 3443, Station D,
Ottawa, ON. KIP 6P4, Canada. E-mail: randerson@mus-nature.ca
Key words. - Weevil; biogeography; inventory; West Indies
The genus Sicoderus is well-represented in the is-
lands of the West Indies with ten recognized species
in two species groups found there (Vanin 1986). The
four members assigned to the S. delauneyi group are
found on the islands of the Lesser Antilles as far north
as Guadeloupe, whereas the six species of the S. tina-
mus group are known from southern Florida and the
Bahamas, Cuba and Hispaniola. Recent collections of
Sicoderus from the British and U.S. Virgin Islands
(and Puerto Rico) bridge the geographical gap be-
tween the ranges of species in these two groups and
represent three undescribed species in the S. tinamus
species group.
Descriptions of these new species follow the format
used in Vanin (1986) to facilitate comparisons with
species treated therein. A revised key to all Sicoderus
species occuring in southern Florida, the Bahamas
and the West Indies is presented.
Specimens are deposited in the Canadian Museum
of Nature, Ottawa, Ontario, Canada (cmnc), the col-
lection of Charles W. O'Brien, Tallahassee, FL.
(cwob), the collection of Michael A. Ivie, Bozeman,
MT. (maic), and the United States National Muse-
um, Washington D.C. (usnm).
Sicoderus ivieorum sp. n.
(figs. 1-5)
Description
Length male, 5.2-6.2 mm; female, 4.8-7.2 mm. In-
tegument black, shining. Rostrum 0. 97-1. 00 X length
elytra in male; 1.09-1.17X length elytra in female.
Antennal insertion submedial in male and female. An-
tenna with article II of funicle 1.49-1.57X length ar-
ticle III in male; 1. 27-1. 29 X length article III in fe-
male. Prothorax with length 1.17-1. 19 X width in
male; 1. 23-1. 26 X width in female. Prothorax con-
stricted anteriorly, globose, widest at anterior one-
third; punctures well-separated, sparse, very fine and
shallow on disk, slightly deeper and larger in anterior
one-quarter and very slightly deeper on flanks; with
scattered, erect setae along anterior constriction. Ely-
tra with length 1. 86-1. 88 X width in male; 1.88-
1.93X width in female; in dorsal view distinctly
widest at midlength, lateral margins convergent both
anteriorly and posteriorly; humeri fully reduced, not
at all angulate; with isolated, erect but short (perhaps
abraded or broken), setae in suturai region; apices con-
jointly rounded. Strial punctures evident, shallow,
moderately fine. Membranous wings lacking. Meta-
sternum with punctures well-separated, moderately
fine and shallow; first row of punctures alongside met-
episternum consisting of 3 or 4 widely spaced, shallow
punctures. Abdominal ventrite I of male moderately
raised in middle near apical margin forming a moder-
ately elevated tubercle with two small subcontiguous
patches of setosity at tip; of female, moderately swol-
len in middle near apical margin but not tuberculate,
not setose. Ventrite V of male with rounded shallow
impression in apical three-quarters; of female, uni-
formly convex. Legs with all femora with distinct ven-
tral tooth; tooth slightly larger on anterior legs; tarsal
claws with small basal tooth. Aedeagus with shallow
apical notch; slightly expanded laterally near apex; in-
129
Tijdschrift voor Entomologie, volume mi, 1998
Figs. 1-2. Sicoderus ivieorum sp. n., male - 1, lateral habitus; 2, dorsal habitus.
ternal sac with complex of denticles and sclerites. Fe-
male not dissected.
Type material. - Holotype male, labelled: 'virgin
is: St. John / Est. CaneelBay, Caneel / Hill Trail fr.
Center / line Rd. 580-600ft / 02nov1992, M.A. Ivie'
(usnm).- Allotype female (usnm) labelled as holo-
type. - Paratypes: 7 males, 3 females. - virgin is-
lands: St. John Est., Caneel Bay, Caneel Hill Trail fr.
Center line Rd. 580-600ft., 2 nov 1992, M. A. Ivie
(1 â , maic); St. John Est. Lameshur Bay, Europa-Lt.
Lameshur Bay, 25 July 1994, M. S.Becker, at night
(1 9, cmnc); St. John Est., Lameshur Bay, Reef Bay
Tr. Europa- Lameshur Bays, 25 July 1994 M.S. Beck-
er, night beating (lo*, cmnc); St. John Est., Ham-
mer-Farm, Cinnamon Bay Trail, 17 julyl994, M.S.
Becker, beating veg (19, maic); St. John Est.,
Molledahl, 15 July 1994, Petroglyph Trail, leaf litter
nr. fresh water, M. S. Becker (lo\ cmnc); St. John
Est., Hope, Reef Bay Trail, 20 July 1994, M. S. Beck-
er coir, beating vegetation (1 S , maic). - british vir-
gin islands: Jost Van Dyke, White Bay, 24 July
1994, M. A. Ivie, beating dead catch-n-keep vines
(2cT, cmnc, maic); Tortola, Windy Hill, 24-29 dec
1993, 300-500', thorn-scrub for., T.K.Phillips coir.
(IS , 19, cmnc, maic).
A single female specimen labelled 'puerto rico:
Guanica For. Res. 26sepl987, MA. Ivie coir. 33m,
beating, thorn scrub' appears to represent this same
species. It is not included in the type series as males
need to be examined to confirm conspecificiry. As
Puerto Rico and St. John are part of the Puerto Rican
Bank (an old continental plate) and are not separated
by deep water, such a distribution would not be sur-
prising.
Derivation of specific name
This species is named for Michael and Donna Ivie
who have worked extensively on increasing our know-
ledge of West Indian, especially Virgin Islands, insects.
Comments
Sicoderus ivieorum can be distinguished from other
Sicoderus in the West Indies by the following combi-
nation of character states: presence of distinct femoral
teeth; elytra with humeri fully reduced and not at all
angulate; sparse and fine punctures of the pronotal
disk, not forming striolae; moderately large and dis-
tinct punctures of abdominal ventrites; moderately
developed tubercle of ventri te I of male; shallow, fine-
ly setose impression on ventrite V of male; and by the
structure of the aedeagus.
130
Anderson: New Virgin Islands Sicoderus
Figs. 3-5.
Sicoderus ivieorum sp. n. male.
— 3, legs, a, front, b, middle, c,
hind; 4, abdomen; 5, aedeagus,
a, dorsal view, b, lateral view.
(3-4, scale line = 1.0mm; 5,
scale line = 0.5mm)
Sicoderus hirsutiventris sp. n.
(figs. 6-10)
Description
Length male, 4.3-6.5 mm. Integument black, shin-
ing. Rostrum 0.80-0.85 X length elytra in male. An-
tennal insertion at apical two-fifths in male. Antenna
with article II of funkle 1. 29-1. 32 X length article III
in male. Prothorax with length 1.17-1.27X width in
male. Prothorax constricted anteriorly, globose,
widest at anterior one-third; punctures well-separat-
ed, very sparse, very fine and shallow on disk, slightly
deeper and larger in anterior one-quarter and slightly
deeper on flanks; with scattered, appressed setae along
anterior constriction. Elytra with length 1. 97-1. 98 X
width in male; in dorsal view distinctly widest at mi-
dlength, lateral margins convergent both anteriorly
and posteriorly; humeri fully reduced, not at all angu-
late; with isolated, erect but short (perhaps abraded or
broken), scattered setae; apices conjointly rounded.
Strial punctures evident, shallow, moderately fine.
Membranous wings present, slightly shorter than ely-
tra in length. Metasternum with punctures indistinct,
shallow and irregular; first row of punctures alongside
metepisternum consisting of 6 widely spaced, very
shallow punctures. Abdominal ventrite I of male
raised in middle near apical margin forming a
markedly elevated tubercle with two small subcon-
tiguous patches of setosity at tip. Ventrite V of male
with large rounded deep, densely setose impression
about as large as entire ventrite V. Legs with front
femora with small, blunt ventral tooth; middle and
hind femora lacking tooth; tarsal claws with small
basal tooth. Aedeagus with shallow apical notch; in-
ternal sac with complex of denticles and sclerites.
Female not known.
Type material. — Holotype male, labelled: 'st.
thomas,vir.is.,/ Ridge above/ CharlotteAmalie/ July
17, 1979', 'Collector:/ C. W. O'Brien' (cwob). - Pa-
ratype, 1 male: virgin islands: St. John Est., Gt.
Cinnamon Bay, Cinnamon Bay Trail, 17 July 1994,
beating, M.S.Becker coir. (1 6 , maic).
131
Tijdschrift voor Entomologie, volume hi, 1998
Figs. 6-10.
Sicoderus h irsutiventris sp. n.,
male — 6, lateral habitus; 7,
dorsal habitus; 8, legs, a,
front, b, middle, c, hind; 9,
abdomen; 10, aedeagus, a,
dorsal view, b, lateral view.
(8-9, scale line =1.0 mm;
10, scale line = 0.5 mm).
132
Anderson: New Virgin Islands Sicoderus
Figs. 1 1-12. Sicoderus vaninisp. n., female— 1 1, lateral habt
tus; 12, dorsal habitus.
Derivation of specific name
This species is named from the Latin 'hirsutus'
meaning hairy and 'venter' meaning underside; in ref-
erence to the large, setose impression of ventrite V in
the male of this species.
Comments
Sicoderus hirsutiventris can be distinguished from
other Sicoderus in the West Indies by the following
combination of character states: presence of a small
tooth on only the front femora; elytra with humeri
fully reduced and not at all angulate; sparse and fine
punctures of the pronotal disk, not forming striolae;
moderately large and distinct punctures of abdominal
ventrites; moderately developed tubercle of ventrite I
of male; large, densely setose impression on ventrite V
of male; and by the structure of the aedeagus.
Sicoderus vanirti sp. n.
(figs. 11-13)
Description
Length female, 5.0-6.6 mm. Integument black,
shining. Rostrum 1.2 1- 1.48 X length elytra in fe-
male. Antennal insertion submedial in female. Anten-
na with article II of funicle 1. 22-1. 33 X length article
III in female. Prothorax with length 1. 22-1. 33 X
width in female. Prothorax constricted anteriorly,
globose, widest at midlength; punctures dense, large
and deep, coalescent, forming striolae on disk, slight-
ly shallower and smaller in anterior one-fifth and to-
wards posterior margin, slightly deeper and larger on
flanks where striolae more distinct; erect setae lack-
ing. Elytra with length 1.78-2. 00 X width in female;
in dorsal view widest at about midlength, lateral mar-
gins slightly convergent both anteriorly and posteri-
133
Tijdschrift voor Entomologie, volume 141, 1998
Fig. 13.
Sicoderus vanini sp. n., fe-
male - legs, a, front, b, mid-
dle, c, hind, (scale line =1.0
mm).
orly; humeri fully reduced, not at all angulate; lacking
erect setae (perhaps abraded or broken); apices con-
jointly rounded. Strial punctures evident, shallow,
moderately fine. Membranous wings lacking. Meta-
sternum with punctures large and deep; first row of
punctures alongside metepisternum consisting of 4-5
closely spaced, large deep punctures. Abdominal ven-
trite I of female markedly convex. Ventrite V of fe-
male uniformly slightly convex. Legs with all femora
lacking tooth; tarsal claws with small basal tooth.
Male not known.
Type material. - Holotype female, labelled: 'British
Virgin Islands/ Tortola, Smuggler's/ Cove 25/
29.viii.1988/ J.LaSalle coll. (ypt)' (cwob). - Paratype,
1 female: british virgin islands: Tortola, Smug-
gler's Cove, 25-29 aug 1988, J.LaSalle coll., yellow
pan trap (1 ? , cwob).
Derivation of specific name
This species is named for Sergio Vanin in honour
of his past work on the Erodiscini.
Comments
Sicoderus vanini can be distinguished from other
Sicoderus in the West Indies by the following combi-
nation of character states: lack of femoral teeth; elytra
with humeri fully reduced and not at all angulate;
dense and deep punctures of the pronotal disk, form-
ing distinct striolae; and large and distinct punctures
of the abdominal ventrites.
Key to Sicoderus adults in West Indies
(including Florida and Bahamas)
1. Tarsal claws with small basal tooth. Florida, Ba-
hamas, Greater Antilles, Lesser Antilles south to
Guadeloupe 2
- Tarsal claws simple, lacking basal tooth. Lesser
Antilles south of Guadeloupe 11*
2. Elytra in lateral view with apex abruptly truncate
(see Vanin 1986: Fig. 315). Hispaniola
S. truncatipennis Vanin
- Elytra in lateral view with apex uniformly ta-
pered. Florida, Bahamas, Greater Antilles, Lesser
Antilles south to Guadeloupe 3
3. Humeri angulate. Florida, Bahamas, Cuba, His-
paniola 4
- Humeri rounded, not at all angulate (Figs. 2, 7,
12). Hispaniola, Puerto Rico, Virgin Islands,
Guadeloupe 7
4. Legs with all femora with distinct (although small
on middle and hind legs) tooth on inner margin. 5
- Legs with only front femora with distinct (al-
though small) tooth on inner margin 6
5. Pronotum on flanks with at least some punctures
coalescent, forming lateral striolae. Aedeagus as
figured in Vanin 1986; Fig. 317. Dominican Re-
public S. championi 'Vanin
- Pronotum on flanks with punctures not coales-
cent, not forming striolae. Aedeagus as figured in
Vanin 1986; Fig. 316. Southern Florida, Ba-
hamas S. tinamus (LeConte)
134
Anderson: New Virgin Islands Sicoderus
6. Elytra usually with a few setae present (see Vanin
1986; Figs. 322-323). Antennal funicle with arti-
cle II 1.1-1.3X length of article III. Aedeagus as
figured in Vanin 1986; Fig. 324. Cuba
S. sleeperi Vanin*
- Elytra lacking setae (see Vanin 1986; Figs. 320-
321). Antennal funicle with article II 1.4-1.5X
length of article III. Aedeagus as figured in Vanin
1986; Fig. 325. Dominican Republic
S. schoenherri\ ranin
7. Ventrites with shallow, small, indistinct punc-
tures; male with tubercle on ventrite I set equidis-
tant between posterior margin of hind coxal cavi-
ty and posterior margin of ventrite I. Guadeloupe
S. delauneyi (Chevrolat)
- Ventrites with at least anterolateral portions of
ventrite I and ventrite II along suture with ven-
trite I with moderately deep, moderately large to
large, distinct punctures; male with tubercle on
ventrite I set immediately anterior to posterior
margin of ventrite I. Hispaniola, Puerto Rico,
Virgin Islands
8. Pronotal disk with punctures large, deep, dense,
coalescent and forming striolae; striolae deeper
and more distinct on flanks (Figs. 11-12). Female
rostrum long (1.21-1.48X length elytra) (Fig.
1 1). British Virgin Islands (Tortola)
S. vanirti sp. n.
- Pronotal disk with punctures very fine and shal-
low, widely spaced, not forming striolae (Figs. 1-
2, 6-7). Female rostrum shorter (1.1 1-1.23X
length elytra; female S. hirsutiventris not known) .
Hispaniola, U.S. Virgin Islands, British Virgin Is-
lands 9
9. Pronotum on flanks with punctures moderately
large and deep, closely separated, coalescent on
some specimens and forming irregular striolae.
Hispaniola S. ramosi (Sleeper)
- Pronotum on flanks with punctures very fine,
shallow, widely separated, not coalescent (Figs. 1,
6). U.S. Virgin Islands 10
10. Legs with all femora with distinct ventral tooth
(Fig. 3); male rostrum longer (0.97-1.00X length
elytra) (Fig. 1); male with small, shallow, finely
setose impression on ventrite V (Fig. 4). U.S. Vir-
gin Islands (St. John), British Virgin Islands (Jost
Van Dyke, Tortola) S. ivieorum sp. n.
- Legs with only front femora with small ventral
tooth (Fig. 8a); male rostrum shorter (0.80-
0.85 X length elytra) (Fig. 6); male with very
large, deep, densely setose impression on ventrite
V (Fig. 9). U.S. Virgin Islands (St. John, St.
Thomas) S. hirsutiventris sp. n.
1 1. Tubercles of ventrite I in male nearly contiguous
(see Vanin 1986; Fig. 302). St. Vincent
S. contiguous Vanin*
- Tubercles of ventrite I in male not contiguous
(see Vanin 1986; Figs. 300-301) 12
12. Tubercles of ventrite I in male separated by a dis-
tance nearly equal to the width of the abdominal
process at base (see Vanin 1986; Fig. 300). St.
Vincent S. remotus Vanin*
- Tubercles of ventrite I in male separated by a dis-
tance varying from one-quarter to one-third of
the width of the abdominal process at base (see
Vanin 1986; Fig. 301). Grenadines, Grenada,
Trinidad and Tobago S. propinquusVan'm*
8 * Species not seen by the author.
Acknowledgements
Thanks are due to Michael and Donna Ivie for
inviting me to become involved in the Virgin Islands
Beetle Project and for lending some of the specimens
on which this paper is based. I also thank Charles W.
O'Brien for the loan of specimens and for review of
the manuscript. Susan Laurie-Bourque prepared the
line illustrations. This paper is a contribution to the
NSF sponsored Virgin Islands Beetle Project, Michael
A. Ivie (Principal Investigator).
Literature cited
Vanin, S.A. 1986. Systematics, cladistic analysis, and geo-
graphical distribution of the tribe Erodiscini (Coleoptera,
Curculionidae, Otidocephalinae). - Revista Brasileira de
Entomologia 30: 427-674.
Received: 1 5 September 1 998
Accepted: 15 December 1998
135
Tijdschrift voor Entomologie, volume 141, 1998
BOOK REVIEW
F. Bos & M. Wasscher, 1997. Veldgids libellen. - Stichting
Uitgeverij KNNV, Utrecht. NLG 49.95 p/p excl. Available
from KNNV Uitgeverij, Oudegracht 237, 35 1 1 NK Utrecht,
The Netherlands. [Field guide to Odonata, In Dutch]
The study of the distribution and status of dragon-
flies (Odonata) has become very successful among
amateur entomologists of western Europe, and the re-
sults are equally impressive. Study groups are investi-
gating nearly every kilometre square for dragonflies,
resulting in overviews of the distribution of the
species on local, regional or national scale. These ac-
tivities are particularly welcome, since the distribu-
tion of many species appears to be highly dynamic at
present. The range expansion of species as Erythrom-
ma viridulum, Crocothemis erythraea and Anax parthe-
nope during the last twenty years has been carefully
documented. On the other hand, also rare and threat-
ened species have been mapped and many formerly
unknown populations have been discovered, provid-
ing the necessary data for measures to protect bio-
topes optimal for Odonata.
Of course, the basic data of maps shall be reliable. It
is impossible and also undesirable, that such atlases are
only based on material in collections. Besides, in many
parts of Europe some, and in some countries all
species are protected by law, making even the effort to
collect them unlawful. These facts are hints for the ob-
vious need for field guides to dragonflies, as there are
so many to birds, mammals and butterflies. Various of
such guides have been published during the last few
years. The concept of some of these guides is highly
original. Three authors, Tieneke de Groot, Weia
Reinboud and Marcel Wasscher prepared (in Dutch)
the 'Odon-key for identification of dragonflies with-
out collecting' (published in 1993 by the Jeugd-
bondsuitgeverij, Donkerstraat 17, 3511 AR Utrecht),
aiming at young amateurs in The Netherlands and
Belgium. This simply produced key is based on a pro-
found field knowledge of the species, leading the start-
ing and more advanced odonatist through the identi-
fication process in an authoritative way. The
illustrations are simple, but adequate. Based on a com-
pletely different concept, but equally innovative, is
Steve Brooks' (editor) 'Field guide to the dragonflies
and damselflies of Great Britain and Ireland'. All
species are represented by several colour illustrations.
The texts are well-structured, with descriptions divid-
ed into paragraphs on jizz, field characters and similar
species.
The 'Veldgids libellen' (Field guide to Odonata) by
Frank Bos and Marcel Wasscher is a more conven-
tional, but attractively produced guide to the species
of western, northern and central Europe. More than
160 colour photographs are included. The common
species are represented by both adult male and fe-
male, the less common only by male or female. Pages
opposite to photos have texts on recognition, flight
period, habitat, range and distribution, including
small distribution maps for Europe, and an indication
in colour of how common a species is in The Nether-
lands. Apart from the scientific name, also the Dutch,
English, German, French and Swedish names are
provided. Also, there are the usual introductory chap-
ters on morphology, life cycle and habitats.
The quality of most colour illustrations is satisfac-
tory, but really good photographs are scarce. The
texts, although obviously written by experts, suffer
somewhat from a lack in format. Beginners will meet
difficulties in finding the correct name for a speci-
men, particularly so, since an identification key is
missing. Illustrations of structural details are given in
some cases only, e.g. dorsal views of the prothorax of
Coenagrion species.
The first edition included several errors, which
were corrected in the second edition of December
1998. The main addition is the inclusion of the Red
Data Lists of The Netherlands and Flanders (Bel-
gium). The printing of some photographs has been
improved, and some photographs have been replaced
by better ones. The best news of this reprint is that
more than 4000 copies of a colour guide to the drag-
onflies written in Dutch were sold in less than one
year, indicating how popular the study of dragonflies
nowadays is.
[J. van Tol]
136
Ping-Ping CHEN & Herbert ZETTEL
Naturhistorisches Museum Wien, Austria
A TAXONOMIC REVISION OF THE ORIENTAL
WATER STRIDER GENUS VENTIDIUS DISTANT
(HEMIPTERA, GERROMORPHA, GERRIDAE)
Chen, P. P. & H. Zettel, 1999. A taxonomie revision of the oriental water strider genus Ventid-
ius Distant (Hemiptera, Gerromorpha, Gerridae). - Tijdschrift voor Entomologie 141 [1998]:
137-208, figs. 1-266, maps 1-6, tables 1-3. [issn 0040-7496]. Published 1 March 1999.
The Oriental water strider genus Ventidius Distant is redescribed and revised. The morpholog-
ical characters of Ventidiopsis Miyamoto and Esakia Lundblad and their relationships with Ven-
tidius are discussed. Ventidiopsis Miyamoto is ranked as a subgenus of Ventidius (stat. n.). Six
species are described as new: Ventidius (s. str.) longitarsus sp. n. from Viet Nam, V. (s. str.) pi-
losus sp. n. from Indonesia (Nusa Tenggara Timur, Nusa Tenggara Barat), V. (s. str.) polhemo-
rum sp. n. from Malaysia (Sabah, Sarawak), V. (Ventidioides) heissi sp. n. from Malaysia
(Sarawak), V. {Ventidioides) nieseri sp. n. from Brunei, Malaysia (Sarawak) and Indonesia (Kali-
mantan), and V. {Ventidiopsis) yangae sp. n. from Malaysia (Sabah). The following species are
redescribed: Ventidius (s. str.) aquarius Distant, V. (s. str.) usingeri Hungerford & Matsuda, V.
(s. str.) harrisoni Cheng, V. (s. str.) malayensis Hungerford & Matsuda, V. (s. str.) henryi Esaki,
V. (s. str.) hungerfordi Cheng, V. (s. str.) werneri Hungerford & Matsuda, V (s. str.) modulatus
Lundblad, V { Ventidioides) kuiterti Hungerford &C Matsuda, V { Ventidioides) karen Lansbury,
V. {Ventidioides) pulai Cheng, V. {Ventidioides) lundbladi Miyamoto, V. {Ventidioides)
xiphibion Chen & Nieser, V. { Ventidioides) xyele Chen & Nieser. V. { Ventidioides) kurtokalami
Chen Sc Nieser, and V. {Ventidiopsis) imadatei Miyamoto. Type specimens of most species were
re-examined, except those of V. (s. str.) distanti Paiva, V. (s. str.) sushmae Gupta, and V. lund-
bladi, which were unavailable. The following taxa are considered as synonyms (junior syn-
onyms in brackets): Ventidius (s. str.) hungerfordi Cheng (= Ventidius wallacei Lansbury, syn.
n.), V. (s. str.) modulatus Lundblad (= V. pubescens Cheng, syn. n., = V. c hinai H ungerford &
Matsuda, syn. n.). V sushmae Gupta is regarded as a nomen inquirendum. One new combina-
tion is established: Ventidius { Ventidiopsis) imadatei Miyamoto, comb. n. The identity of V. dis-
tanti is discussed. A lectotype of V. modulatus Lundblad is designated.
The species are arranged in three subgenera and five species groups: Ventidius (s. str.): V. aquar-
z'ztf-group, V. modulatus-gioup; Ventidius {Ventidioides): V. kuiterti-group, V xiphibion-group;
Ventidius { Ventidiopsis): V. imadatei-gioup. Descriptive notes and illustrations of pertinent char-
acters are presented for all species. Tables of measurements and an identification key for apter-
ous males of all described species are presented.
Correspondence: Pingping Chen, c/o Naturhistorisches Museum Wien, Zweite Zoologische
Abteilung, A-1014 Wien, Austria.
Key words. - Gerridae; Ventidius; revision; new species; distribution; identification key; Orien-
tal region.
Contents Subgenus Ventidius Distant 144
The Ventidius aquarius-gvoup 148
Introduction 137 The Ventidius modulatus-gxoup 157
Notes on relationships 139 Subgenus Ventidius (Ventidioides) Hungerford &
Checklist 139 Matsuda 176
Material and methods 140 The Ventidius kuiterti-group 178
List of depositories 140 The Ventidius xiphibion-gwup 188
Acknowledgements 141 Subgenus Ventidius (Ventidiopsis) Miyamoto ... 199
Taxonomy 14 1 The Ventidius imadatei-gxoxxp 199
Genus Ventidius Distant 141 References 204
Key to the subgenera of Ventidius 144 Tables 206
137
Tijdschrift voor Entomologie, volume mi, 1998
Introduction
This revision deals with the species of the Oriental
water strider genus Ventidius Distant, 1910, which
was last revised by Hungerford & Matsuda (I960).
Ventidius was established by Distant for a Halobatine
gerrid from 'Travancore, India', which he named
Ventidius aquarius (Distant 1910a: 150). In the same
year, Distant (1910b: 157) figured both macropter-
ous and apterous specimens of the same species.
Bergroth (1911) considered Ventidius a synonym
of Metrocoris Mayr, which was corrected by Esaki
(1929). Paiva (1918) described V. distanti from
'Yawnghwe State' (today belonging to Myanmar), a
species which was misinterpreted several times by the
following authors: Dover (1929), Esaki (1930),
Hungerford & Matsuda (1960). Esaki (1928) and
Lundblad (1933) described two further species, V.
henryi Esaki from Sri Lanka and V. modulatus Lund-
blad from Java.
Hungerford & Matsuda (I960) revised the genus,
recognizing nine species, including five new species
from Myanmar (Burma), Malaysia, and the Philip-
pines. Beside giving excellent pictures of the dorsal
view of all the nine species, they also presented figures
for characters of the new species ( V. malayensis, V. min-
gevi, V. werneri, V chinai, and V. kuitertt). They also
divided the genus into two subgenera: Ventidius (s. str.)
and Ventidioides Hungerford & Matsuda, 1960 (type
species: V. kuitertt) according to four characters: shape
of posterolateral angle of metacetabula bilobate or
transverse; fore femur with or without a tubercle;
mesosternum with or without a tubercle; and para-
meres large or small and asymmetrical or symmetrical.
Matsuda (1960) gave an extensive morphological de-
scription for the genus and set it close to the Mada-
gassian genus Ewymetropsielloides Poisson 1956, and
the Oriental genus Esakia Lundblad 1933, and within
the newly established tribe Metrocorini Matsuda I960.
Cheng (1965) reviewed the Ventidius species of
Malaysia and Singapore and described four species as
new from this area: Ventidius pulai, V. harrisoni, V.
hungerfordi, and V. pubescens.
Since that time, only smaller notes on Ventidius and
sporadic descriptions of new species have appeared.
Miyamoto (1967) added V. lundbladi as a new species
from Thailand, and Ventidius susbmae was described
by Gupta (1981) from India (here regarded as a nomen
inquirendum). Andersen (1982) figured the distribu-
tion of Metrocorini, indicating that its range in the
Malay archipelago is limited by the Wallace's Line.
Later on, several species of Ventidius and Metrocoris
were described from Sulawesi. Lansbury (1990) de-
scribed V. ( Ventidioides) karen from Thailand, and V.
(s. str.) wallacei from Malaysia, the latter a synonym of
V. hungerfordi (syn. n.). Three new species of the sub-
genus Ventidioides, V kurtokalami (from Sabah), V.
xiphibion, and V. xyele (both from Sulawesi), were de-
scribed by Chen & Nieser (1992), increasing the num-
ber of species to 20. Kovac & Yang (1989) and Yang &
Kovac (1995) reported new records of V. malayensis, V
harrisoni, V. hungerfordi, V. pulai and V. modulatus
from the Malaysian Provinces of Pahang and Perak, re-
spectively. Finally, Thirumalai (1996) reported V.
( Ventidioides) kuiterti as a new record from NE India.
The genus Ventidiopsis was erected by Miyamoto
(1967) for the single species Ventidiopsis imadatei
Miyamoto 1967, from Borneo. Because of close rela-
tionship between Ventidiopsis and Ventidioides, both
taxa are regarded as subgenera of Ventidius in the pre-
sent study.
Since the genus Esakia, which here is regarded as
the genus most closely related to Ventidius, was de-
scribed by Lundblad (1933), six species were described
from Myanmar (Burma), Malaysia, Borneo (Brunei),
and the Philippines (Luzon) (Cheng 1966, Hun-
gerford & Matsuda 1958, Miyamoto 1967). Further
new species are known to the authors from Viet Nam,
Borneo, and the Philippines (Palawan). In the present
study, Esakia ventidioides Lundblad 1933, from
Sumatra (Lundblad 1933) was checked.
The Madagassian genus Eurymetropsielloides, which
is according to Matsuda (1960) the sister taxon of
Ventidius+ Esakia, differs from both of these genera by
the metacetabula having strongly downcurved pos-
terolateral corners (plesiomorphic character) and an
unusual colour pattern (apomorphic character) (see
also Matsuda I960).
In the present taxonomie revision we recognize 24
species of Ventidius, including six new species de-
scribed from Viet Nam, Borneo and Indonesia (Sum-
ba, Sumbawa). They are arranged in three subgenera
and five species-groups (see following check-list) .
Ventidius species are small, black and green (turns
to yellowish after death) water striders which are typi-
cal inhabitants of slow-flowing forest streams in South
and Southeast Asia. Their preferred habitats are-as far
as we know for a few species-lotic areas of brooks,
streams and rivers, where they stroke and glide across
the surface. Because of their preference for the slow-
flowing streamlets, Ventidius are mainly encountered
in hilly and lowland areas. The life history of Ventid-
ius species is unknown. Murphy (1990) described the
habitat in Singapore of V. (s. str.) hungerfordi Cheng
as follows: 'The outflow stream of the Nee Soon
swamp forest is small but fairly recent alluvia, and is
deeply shaded. Here the open water is populated by
Ventidius harrisoni and Rheumatogonus intermedius,
sandbanks by Amemboa riparia and Tenagogonus
quiquemaculata and under the overhanging root mats
the tiny haloveliinae Strongilovelia occurs. [...] Farther
upstream a network of shallow channels on peat but
138
Chen & Zettel: Revision ofVentidius
still with moving water is populated by Metrocoris
tenuicornis, stiller water with leaf beds by Tenagogonus
insularis. Throughout the whole system, leafy banks
carry large numbers of Rhagovelia femorata. Elsewhere
in the swamp forest of Nee Soon are larger but now in-
accessible rivers where Ventidius hungerfordi, Esakia
fernanndoi and Cylindrostethus costalis were collected.'
Notes on relationships
In this paragraph, relationships between the taxa
Ventidius, Ventidioides, Ventidiopsis and Esakia are
discussed.
The subgenus Ventidius s. str. consists of two rather
different species-groups of closely related species
which are both widely distributed in the Oriental re-
gion. Although both of these groups are clearly de-
fined by several apomorphic characters, the only
synapomorphy of both groups may be the reduction
of paramere size. It is not proved, if the subgenus Ven-
tidius s. str. is a monophyletic taxon or a paraphyletic
group at the base of the genus Ventidius. In the latter
case, the species of the V. aquarius-group belong to
Ventidius s. str., and the V. modulatus-group should be
raised to a presently unnamed subgenus.
Ventidioides forms a rather nonhomogeneous accu-
mulation of species which are divided here into two
established species-groups; all characters discussed by
Hungerford & Matsuda (1960), Matsuda (I960),
Cheng (1965), and Lansbury (1990) refer to species of
the V. kuiterti-group, which inhabits the Southeast
Asian mainland. The V xiphibion-gwup contains five
species from Borneo and Sulawesi. The only synapo-
morphy for all Ventidioides species is the presence of
special structures on mesosternum of males (tubercle
with setae or row of setae). All other 'apomorphic'
characters mentioned by Hungerford & Matsuda
(1960) and Matsuda (1960) are either not developed
in all species (tubercle of fore femur lacking in the V.
xiphibion-group) or are also developed in Ventidiopsis.
If Ventidioides proves to be polyphyletic, then the V.
xiphibion-group should be raised to a presently un-
named subgenus.
When Miyamoto (1967) described Ventidiopsis
imadatei, only the female was known, which has strik-
ing characters, and therefore this species was placed in
a new genus different from Ventidius. After studying
the male of V. imadatei, we found some reasons to in-
clude Ventidiopsis as a subgenus within Ventidius and
to treat it within this revision. At least two striking
characters are regarded as synapomorphies of the sub-
genera Ventidioides and Ventidiopsis stat. n., i.e. the
bilobate posterolateral angle of metacetabula and the
asymmetrical parameres.
The species of Esakia form a homogeneous group
with several synapomorphic characters, e.g. the dilat-
ed third antennal segment of male, the reduced fore
wing venation, and a flatter, anteriorly more widened
body (see also Matsuda [I960]). But the complete ab-
sence of an anterolateral separation of the metanotum
(which is present in Esakia by a longitudinal eleva-
tion) is the only character which we found useful as
synapomorphy of the proposed monophylum Ven-
tidius+ [ Ventidioides+ Ventidiopsis] .
The monophyletic status of Ventidius+ Esakia is
supported by several characters, e.g. short pronotum,
larger eyes, and rather strongly flattened metacetabula
(Matsuda 1960), which is confirmed by this study.
This provisional arrangement of genera and subgenera
should be proven in a subsequent cladistic analysis.
An analysis of the present distribution of Ventidius
suggests that Malaysia has the richest biodiversity.
Checklist
Ventidius Distant 1910
Subgenus Ventidius s. str.
Ventidius aquarius-group
1. aquarius Distant 1910
2. usingeri Hungerford & Matsuda, 1960
3. harrisoni Cheng, 1965
4. malayensis Hungerford & Matsuda, 1960
5. longitarsus sp. n.
Ventidius modulatus-group
6. henryi Esaki, 1928
7. hungerfordi Cheng, 1965
= wallacei Lansbury, 1990
8. werneri Hungerford & Matsuda, 1960
9. polhemorum sp. n.
10. pilosus sp. n.
11. modulatus Lundblad, 1933
= chinai Hungerford & Matsuda, 1960
= pubescens Cheng, 1965
12. ? distanti Paiva, 1918
13. ? sushmae Gupta, 1981
Subgenus Ventidioides Hungerford & Matsuda, I960
Ventidius kuiterti-group
14. kuiterti Hungerford & Matsuda, I960
15. karen Lansbury, 1990
1 6. pulai Cheng, 1 965
17. lundbladi Miyamoto, 1967
Ventidius xiphibion-group
18. xiphibion Chen & Nieser, 1992
19. xyele Cnen & Nieser, 1992
20. kurtokalami Chen & Nieser, 1992
21. nieseri sp. n.
22. heissi sp. n.
Subgenus Ventidiopsis Miyamoto, 1967
Ventidius imadatei-group
23. imadatei Miyamoto, 1967
24. yangae sp. n.
139
Tijdschrift voor Entomologie, volume i4i, 1998
Figs. 1-5. Diagrams of Ventidius sp. with lines indicating measurements. - 1, apterous male (appendages removed) in dorsal view
(a, interocular width; b, eye width; c, posterior eye width; d, head width; e, pronotum width; f, median length of body; g, body
width across acetabula); 2, macropterous form (appendages removed) in dorsal view (h, median length of pronotum; i, humeral
width of pronotum; j, length of lateral margin from anterior angle to humerus of pronotum; k, length of lateral margin from
humerus to apex of pronotum; 1, length of fore wing from humerus to apex); 3, lateral view of genital segments (py, pygophore;
pr, proctiger; pa, paramere; en, endosoma); 4, dorsolateral view of endosoma (as, apical sclerites; ds, dorsal sclerites; Is, lateral scle-
rites; 2ls, second lateral sclerites; se, ductus seminis; vs, ventral sclerites); 5, lateral view of endosoma (3ls, third lateral sclerites).
Material and methods
The revision is chiefly based on material borrowed
from various museums and private collections (listed
as depositories below). Parts of the material were re-
cently collected by the authors in Eastern Malaysia,
Thailand, and the Philippines. The holotypes of all
new species are deposited in major institutions and
museums as indicated in the taxonomie section.
Olympia optics, including a Camera Lucida, were
used in studying and drawing the specimens. Exami-
nation of the male terminalia is essential for a safe
identification of most species of Ventidius. The male
genital segments were detached from the specimens.
Before dissection, the segments were cleared in 10%
potassium hydroxide for 12-24 hours at room tem-
perature, or till the endosoma became clearly visible.
For studies of thoracic hair layers in species of the V.
modulatus-group a Wild binocular optic with magni-
fication up to 1000X was used.
All measurements in the descriptions are in mil-
limetres. The tables of measurements (table 1-3) give
the means of measurements of six specimens for each
species or, if less than six specimens were available, of
all specimens studied. Interocular width is the short-
est distance between eyes (fig. 2, a); eye width is the
greatest width of the eye measured perpendicular to
the longitudinal axis of the head (fig. 2, b); head
width is the greatest width of head measured across
eyes (fig. 2, d). Width of pronotum in the apterous
form and pronotai measurements in the macropter-
ous form as illustrated in figs. 1 and 2, respectively.
When measuring the antennal segments, the small in-
ternodal piece between segments 2 and 3 was includ-
ed into the length of segment 3. Terms of male geni-
talia see figs. 4-5.
140
Chen & Zettel: Revision ofVentidius
List of depositories
AMNH American Museum of Natural History, New
York, U.S.A.
BiMC B.P. Bishop Museum, Honolulu, U.S.A.
BMNH The Natural History Museum, London,
U.K. (= former British Museum [Natural
History]).
CNHM Chicago Natural History Museum, Chicago,
U.S.A.
FMHF Finnish Museum of Natural History,
Helsinki, Finland.
FMNH Field Museum of Natural History, Chicago,
U.S.A.
GCLB Gupta Collection, Los Banos, Philippines.
JTPC Colorado Entomological Museum,
Englewood, Colorado, U.S.A. (= formerly J.
T. Polhemus Collection).
KKUA Khon Kaen University, Faculty of
Agriculture, Department of Entomology,
Khon Kaen, Thailand.
NCTN Nieser Collection, Tiel, The Netherlands.
NHMW Naturhistorisches Museum Wien, Vienna,
Austria.
OUMC Hope Department, Oxford University
Museum, Oxford, U.K.
ppcc P. P. Chen Collection, Beijing, China.
RMNH Nationaal Natuurhistorisch Museum
Naturalis (formerly Rijksmuseum van
Natuurlijke Historie), Leiden, The
Netherlands,
sccu W. D. Shepard Collection, California State
University, Sacramento, U.S.A.
SEMC Francis Huntington Snow Entomological
Museum, University of Kansas, Lawrence,
Kansas, U.S.A.
SMNH Swedish Museum of Natural History,
Stockholm, Sweden.
SPCM Sabah Parks Collection, Kota Kinabalu,
Malaysia.
TMBC Magyar Termeszettudomanyi Museum (=
Hungarian Natural Science Museum),
Budapest, Hungary.
UMSM Universiti Malaysia Sabah, Kota Kinabalu,
Malaysia.
UPLB University of the Philippines, Los Banos,
Natural History Museum, Philippines.
ZCMG G. Zimmermann Collection, Marburg,
Germany.
ZCWA H. Zettel Collection, Vienna, Austria.
ZMAC Instituut voor Systematiek en
Populatiebiologie (= formerly Zoologisch
Museum), University van Amsterdam,
Amsterdam, The Netherlands.
ZMUC Zoological Museum, University of
Copenhagen, Copenhagen, Denmark.
ZRCS Zoological Reference Collection, Singapore.
Acknowledgements
We are indebted to the following persons who made types
and/or other material for this revision available: N. M. An-
dersen (zmuc), R. W. Brooks (semc), J. Duffels (zmac), E.
Heiss (Innsbruck), I. Lansbury (oumc), P. Lindskog (smnh),
J. Margerison-Knight (bmnh), N. Nieser (nctn), S. Oygur
(amnh), P. P. Parillo (fmnh), D. A. Polhemus (formerly
bimc), J. T. Polhemus (jtpc), R. Schuh (amnh), J. van Tol
(rmnh), T. Vâsârhelyi (tmbc), C. M. Yang (zrcs), and Y.
Hanboonsong (kkua). We thank L. Dembicky and P. Pa-
cholatko (both Brno) for depositing specimens in the nhmw.
We are most grateful to S. Wongsiri, S. Leepitakrat, R.
Thapa, U. Booncham (all in Chulalongkorn University,
Bangkok), S. Piyapichart, Sudarak (both Naresuan Univer-
sity, Phitsanulok), T. Jamjanya, Y. Hanboonsong, N. Sang-
pradub, Dun, Wirote, Nutcharee, Jum (all in Khon Kaen
University), Virode Naknan (Nam Nao NP), and P.
Schwendinger (Innsbruck and Chiang Mai) for their com-
pany and help during our field work in Thailand.
The junior author is deeply indebted to Maryati Mo-
hamed (umsm) and W. Hödl (University of Vienna) who
enabled his field trip to Sabah. Further, his thanks are due to
Homathevi R. (umsm), G. Gunsalem (spcm) and the staffof
the Danum Valley Field Centre for the hospitality during
his stay in the research station. He highly appreciated the
help of the wawa students group (G. Grabenweger, E.
Griinberger, and G. Schindler) who assisted him in the
Danum Valley. Further, the junior author wants to express
his special thanks to J. Recuenco-Adorada, V. P. Gupta, and
A. C. Sumalde (all in uplb) as well as his many other Philip-
pine friends who supported his field work in the Philippines.
We are deeply indebted to J. T. Polhemus for a scientific
and linguistic review of the manuscript, to I. M. Kerzhner
(St. Petersburg) and N. Nieser for discussing the manuscript
and the geographic distribution patterns, and to Ch. Hech-
er (Vienna) for mapping numerous distributional data.
The study was enabled by the stay of the first author in
Vienna, which was financially supported by the Österreich-
ischer Austausch-Dienst (öad).
Taxonomy
Genus Ventidius Distant
Ventidius Distant, 1910: 149-150. Type-species by mono-
typy: Ventidius aquarius Distant, 1910.
Ventidius Distant; Bergroth 1911: 186 (synonymized Ven-
tidius with Metrocoris Mayr); Esaki 1928: 509-511 (de-
scr., new species); Esaki, 1929: 417 (confirmed as valid
genus); Esaki 1930: 18; Hungerford & Matsuda 1960:
323-343 (descr., illustr., key, new species, new subgenus);
Matsuda 1960: 313-316 (morphological descr., syst., il-
lustr.); Cheng 1965: 153-163 (key to species of Malay
Peninsula, descr., illustr., new species); Gupta 1981: 97-
99 (descr., illustr., key to subgenera, new species); Kovac
& Yang 1989: 285 (list, in Pahang, Malaysia); Chen &
Nieser 1992: 156-159 (descr., illustr., new species); Yang
& Kovac 1995: 293 (list, Perak, Malaysia); Zettel &
Chen 1996: 152, 180 (list, Viet Nam).
Nomenclatorial notes. - The genus Ventidius was
established by Distant (1910) for the new species V.
aquarius, known from both apterous and macropter-
141
Tijdschrift voor Entomologie, volume i4i, 1998
Fig. 6. Ventidius (s. str.) usingeru apterous male, length 3.65 mm.
ous forms. Bergroth (1911) considered the genus
Ventidius Distant identical with Metrocoris Mayr,
stating that the distinguishing characters given by
Distant (1910) were only specific characters, and that
therefore, Ventidius could not be generally separated
from Metrocoris. Esaki (1929) pointed out that Ven-
tidius was wrongly synonymized by Bergroth (1911),
and insisted that Ventidius is a very distinct genus.
Since then, no more confusion has been created on
the generic status, which has been accepted by all re-
cent authors.
Redescription
Dimensions. - Males smaller than females, length
of body usually 2.37-4.60 (6), 2.50-4.40 (9); maxi-
mal width of body usually 1.60-3.10 (<?); 1.74-2.96
(9); width of head across eyes 1.01-1.65 (<?), 1.04-
1.61(9).
Colour. - General colouration yellowish to light
brown, overlain with complex black patterning. Sever-
al small brownish marks or one big dark mark present
on interocular space of head, in some species too vague
to see. Inner margin of eyes usually with brownish
142
Chen & Zettel: Revision ofVentidius
stripes. Eyes varying from greyish to black. Base of ros-
trum dark. Antennae dark except basal part of first
segment; in two species segments 3 and 4 with yellow-
ish bands. Anteclypeus, labrum and apical part of labi-
um dark. Pronotum of apterous form with lateral dark
stripes or a T-shaped dark mark, in some species total-
ly dark. Pronotum of macropterous form mainly dark
with two yellowish marks (fig. 9). Mesonotum with
lateral dark stripes, but showing broad variation, often
forming a pale triangular area between the pale stripes
(fig. 6). Metanotum either totally pale or with lateral
dark marks or stripes, posterior margin with a triangu-
lar dark mark, which is confluent with dark abdomi-
nal tergite 1 . In some species the pale marks on metan-
otum confluent with the pale marks on mesonotum
and together forming a big mushroom-shaped or
pear-shaped pale mark (figs. 54, 183). Propleura yel-
lowish, in some species with a dark mark; a dark stripe
usually running through the length of mesopleura;
metacetabula yellowish, usually dark at protruding
posterior angle, in some species more darkened and
with a long yellowish mark in the central part. Narrow
dark mark on anterior half of ventral metacetabula.
Fore femur yellowish to light at basal third or half; fore
tibia either totally dark or pale at base of external sur-
face; fore tarsi totally dark. Middle and hind legs to-
tally dark, in some species the middle legs yellowish
basally. Fore wings dark brown with dark veins (fig.
155). Abdomen mainly dark dorsally, with pale
marks. Ventral surface of body yellowish.
Structural characters. - Body triangular or subtri-
angular in dorsal outline, only moderately dorso-ven-
trally flattened; body clothed with short fine pubes-
cence. Head bluntly projecting anteriorly of eyes,
with broadly rounded anterior margin; head length in
apterous form about 1.5 times as long as pronotum
along median line. Eyes relatively large, narrower
than the width of interoculus; posterior margin in lat-
eral view covering propleura and anterior part of
mesopleura. Vertex along inner margin of eyes bear-
ing 5-6 stiff setae, interocular with 4 pairs of tri-
chobothria at vertex, the third pair shorter than the
other pairs and extending externally, the others point-
ing upwards (fig. 134). Antennae slender, in males
usually stouter than in females; inner surface of an-
tennae usually with fine erect pubescence through
apical half of segment 1 to segment (3 or) 4 (except in
V. aquarius-group with very long antennae in males,
longer setae on internal surface and segment 3 of
males modified by an apical indentation). Anterior
margin of pronotum in apterous form slightly sinuat-
ed or concave, posterior margin sinuated. Pronotum
of macropterous form large, forming an acute poste-
rio-medial angle. Lateral suture of metanotum not
reaching the meso-metanotal suture anteriorly. Pleu-
ra clothed with fine pubescence, which in some
species is longer and more conspicuous. Mesosternal
'groove' conspicuous, in most species represented by
a translucent internal ridge. Centre of mesosternum
in some species with a tuft of tiny bristles, which in
certain cases extends towards the anterior margin of
the mesosternum. Metasternum reduced to a small
triangular sclerite, in some species swollen to a tuber-
cle. Abdomen with some golden pubescence dorsally;
caudal margin of posterior abdominal tergites and
genital segments with longer pilosity in male. Male
abdominal sternite 7 much longer than sternite 6,
broadly concave on apical margin. Female abdominal
sternite 7 strongly developed and not modified dis-
tinctively. Laterotergites usually broad. Legs slender,
usually not modified. Fore femur of males slightly
stronger than in females, its ventral surface usually
smooth, but modified in subgenus Ventidioides: in V,
kuiterti-group with a tooth halfway along its inner
surface length (figs. 148-154); in V. xiphibion-gtoup
sometimes with a broad tubercle (figs. 198-202); fore
femur usually bearing a number of long setae, its in-
ner surface sometimes with dense short dark hair
fringe. Inner surface of fore tibia in both sexes
straight, but in two species ( V. kurtokalami and V.
niesen) with an indentation at 2/5 of its length in
male, the remaining 3/5 with distinct dark hair
fringe; its apical finger-like projection with a grasping
comb in male, which is composed by varied number
of bristles and dense pilosity. Inner surface of tarsi
with a row of modified hairs, which vary according to
species. Middle and hind legs long, middle femur
usually slightly shorter than hind femur; middle and
hind femora with a number of long spines, and on
dorsal surface with one or two long trichobothria-like
setae basally.
Male genital segments usually relatively large. They
comprise the following structures: the cylindrical seg-
ment 8 (s8); the boat-shaped pygophore (fig. 3, py,
segment 9); the plate-shaped proctiger (pr, segment
10+11); the parameres (pa), which are varied in size,
symmetrical and small in subgenus Ventidius, but
asymmetrical and longer in subgenus Ventidioides,
and of high diagnostic importance; and the phallus
(ph). The distal section of the phallus is the endoso-
ma, which is armed with a number of sclerites. In the
descriptions, the endosoma sclerites and other struc-
tures are named as follows (figs. 4, 5): dorsal sclerites
(ds), apical accessory sclerites (as), lateral sclerites (Is,
paired), ventral sclerites (vs, paired); a second pair of
lateral sclerites occurs in all species, a third pair in
some species (lp); the ductus seminis (se) is supported
by the ventral sclerites.
In females so far only a few useful diagnostic char-
acters have been found, which are usually found also
in males (e.g. some group characters, colour pattern,
body shape, pilosity). There are no distinct differ-
143
Tijdschrift voor Entomologie, volume ui, 1998
ences in female terminalia, except in the subgenus Ve-
tidiopsis. Within species groups not all females can be
identified to species level.
Comparative notes. - Ventidius seems to be most
closely related to the genus Esakia. Main differences
are: thorax anteriorly widened and conspicuously flat
in Esakia, but anteriorly constricted and (usually)
domed in Ventidius; antennal segment 3 dilated in
males of Esakia, but usually rounded in Ventidius
(slightly flattened in subgenus Ventidiopsis); in Esakia
fore wing venation reduced, with closed cells in basal
half only, but more developed in Ventidius, with
closed cells reaching distal half; metanotum with an
antero-lateral separation represented by a longitudi-
nal ridge in Esakia, but fused with metapleura in Ven-
tidius.
Distribution (maps 1-6). - Endemic to the Orien-
tal region, from Sri Lanka to the Philippines and In-
donesia (eastward to Sulawesi and Sumba).
Key to the subgenera of Ventidius
1 . Posterolateral angle of metacetabula truncate or
slightly bilobate in caudal view (figs. 29-38, 75-
87), parameres symmetrical, pronotum yellow
with dark lateral stripes and/or triangular dark
mark in middle of hind margin (except: black in
V. henryi) Ventidius s. su.
- Posterolateral angle of metacetabula distinctly
bilobate in dorsal view (figs. 157-166, 203-212),
parameres asymmetrical, pronotum usually black
2
2. Tergi tes 3 and 4 fused in apterous morphs; 9
(apterous): metacetabula and metacoxae each
with a stout, medially directed and hairy proces-
sus (figs. 240-242), metanotum and anterior ter-
gites concave, meso-metanotal suture with or
without two tufts of hairs; 6 : antenna very stout,
segment 1 thicker than fore tibia, segments 2 and
3 slightly flattened (figs. 249, 252) ... Ventidiopsis
- Tergites 3 and 4 not fused; 9 : without medially
directed processes on metacetabula and metacox-
ae, metanotum and anterior tergites slightly con-
vex to flat, rarely slightly concave (in V. nieseri sp.
n.), meso-metanotal suture without tufts of hairs;
S : antenna slender, segment 1 thinner than fore
tibia, segments 2 and 3 not flattened (fig. 142,
143, 145, 147) Ventidioides
Subgenus Ventidius (s. str.) Distant, 1910
Diagnosis. - Colouration comparatively lighter
than in the subgenus Ventidioides (except in V. hen-
ryi). Antennal segment 1 distinctly shorter than seg-
ments 2-4 together. In some species the subdistal part
of segment 3 modified. Posterolateral angle of metac-
etabula not bilobate or only slightly bilobate. Ventral
surface of male fore femur not modified without a
tooth or tubercle. Mesosternum without tubercle or
row of setae. Parameres symmetrical, small.
Distribution (maps 1-3). - Widely distributed
throughout the Oriental region, from Sri Lanka to
the Philippines, Borneo and Sumba. Highest species
diversity in Malaya.
Identifications. - Nearly all characters used in the
following key for apterous males are also present in
macropterous males (which are unknown in some
species) except pronotum not completely black in
macropterous V. henryi. Females are very difficult to
identify, but species-groups can be easily separated by
the legs. The middle and hind femora are very stout
in females (and males) of the species of V. aquarius-
group, but comparatively slender in species of the V.
modulatus-group. As species of the V. aquarius-group
are mostly allopatric, females may be provisionally
identified after their provenance; only V. malayensis
and V. harrisoni are known to occur in the same areas,
but females of these species can be separated by the
length of the fore tarsus (table 1). Furthermore, there
may be an overlapping zone between the ranges of V.
malayensis and V. longitarsus sp. n. As we could not
find reliable characters to differentiate between fe-
males of these species, identifications of single females
from Indochina are doubtful. Females of the V. mod-
ulatus-group are indistinguishable, except V. henryi
with the typical black pronotum, and V. hungerfordi
with a much flatter body than in V. modulatus. Ven-
tidius modulatus is also distinguishable by the shorter
pilosity of the mesopleura, but this character is diffi-
cult to observe and variable. Other species are al-
lopatrically distributed, but because of the occurrence
of V. modulatus in Borneo, females of V. polhemorum
sp. n. or V. modulatus from this island can only be
identified by the slightly longer pilosity of the first
species.
Key to apterous males of the subgenus Ventidius
1 . Antennal segments very long and slender; length
of segment 1 shorter or subequal to segments 2
and 3 together, usually more than 1.2 mm, at
least with 8 long setae which are evenly scattered
all over its length; segments 2 and 3 with erect,
rather long dark hairs; segment 3 with more or
less distinct subapical indentation (figs. 18-22);
middle and hind femora very stout (fig. 6, 53);
segment 8 with conspicuous deep ventral emar-
gination and lateral tufts of black bristles cau-
doventrally (figs. 39-42); parameres very small,
straight or cucumber-shaped (figs. 43-47) (V
aquarius-gmup) 2
- Antennal segments shorter and less slender; length
of segment 1 longer or subequal to segments 2 and
3 together, usually less than 1.2 mm, and usually
144
Chen & Zettel: Revision ofVentidius
Figs. 7-9.
Ventidius aquarius (female,
antennae and legs removed,
length 3,60 mm). — 7, dotsal
view (holotype); 8, ventral
view (holotype, egg shells visi-
ble); 9, pronotum of macro-
pterous form.
Figs. 10-11.
Ventidius malayensis (male,
antennae and legs removed,
length XX mm). — 10, prono-
tum of macropterous form;
1 1 , dorsal view of apterous
male.
with less than 8 setae, most of them in a subbasal
row; segments 2 and 3 with dense erect silvery pi-
losity; segment 3 without subapical indentation
(figs. 60-66); middle and hind femora normal,
slender (fig. 123); segment 8 without distinct ven-
tral emargination and without lateral tufts of
black bristles; parameres slightly longer, in most
species apically widened (figs. 97-101), except in
V. henryi(see fig. 96) {V. modulatus-group) 6
Segment 2 of fore tarsus only moderately long,
surpassing the claws only slightly, fore tibia inter-
nally with a row of stout spinules (fig. 23, 25,
26), hairs on apical portion of antennal segment
3 only slightly longer than the normal erect hairs
3.
all over the length of the segment and shorter
than width of segment at this location (figs. 18,
19) 3
Segment 2 of fore tarsus very long, surpassing the
claws by more than their length (figs. 27, 28),
fore tibia internally without a row of stout spin-
ules, antennal segment 3 apically with conspicu-
ous long hairs which are longer than width of seg-
ment at this location (fig. 22) 5
Antennal segment 3 about 0.6 times as long as
segment 2, straight and with a sharp indentation
subapically, spinules on internal face of fore tibia
distinctly increasing in length from base to tip
and becoming hair-like in distal third, segment 8
145
Tijdschrift voor Entomologie, volume i4i, 1998
Figs. 12-17. Lateral view of apterous form, showing colour pattern of pleura. - 12, Ventidius aquarius (holotype); 13, V. Usin-
gen; 14, V. harrisoni (paratype); 15, V. malayensis; 16-17, V. longitarsus (types).
4.
with broader tuft of hairs ventrocaudally 4
Antenna! segment 3 about 0.8 times as long as
segment 2, with a very shallow indentation sub-
apically (fig. 20); spinules on internal face of fore
tibia only sligthly increasing in length from base
to tip and becoming only slightly thinner towards
apex of tibia (fig. 26), segment 8 with narrow
tufts of hairs ventrocaudally (fig. 41) (Malaya,
Borneo) V. harrisoni
Mesopleural stripe ending at or at short distance
from the second spiracle, ventral incision of seg-
ment 8 broad and less deep (fig. 40), parameres
more straight and with numerous small hairs ven-
trally (fig. 44) (Philippines) V. usingeri
— Mesopleural stripe ending in middle of meso-
pleura in long distance from the second spiracle,
ventral incision of segment 8 narrower and deep-
er (fig. 39), parameres more curved and without
small hairs ventrally (fig. 43) (Sri Lanka, South
India) V. aquarius
5. Segment 2 of fore tarsus very long (fig. 28), an-
tennal segments 2-4 slightly thicker than in the
following species, colouration variable (Viet
Nam) V. longitarsus
- Segment 2 of fore tarsus slightly shorter (fig. 27),
antennal segments 2-4 extremely thin, coloura-
tion always bright (Thailand, Malaya, Borneo) ..
V. malayensis
146
Chen & Zettel: Revision ofVentidius
Figs. 18-22. Dorsal view of male right antennae. - 18, Ventidius aquarius; 19, V. usingeri; 20, V. harrisoni (paratype); 21. V.
malayensis; 22, V. longi tarsus (holotype).
6. Pronotum completely black (fig. 54); distal part
of parameres parallel-sided (fig. 96); thorax
clothed with dense, long pubescence and fore fe-
mur slightly flattened (Sri Lanka, South India) ..
V. henryi
— Pronotum completely or mainly yellowish, or at
least with a large yellowish mark in middle; distal
part of parameres club-shaped or at least slightly
dilated apically (figs. 102-113); either thorax
without dense, long pubescence or fore femur not
slightly flattened 7
7. Antennal segment 1 very slender, 12.5-15 times
as long as maximum width; thorax with less dense
pubescence, sometimes relatively dense in dorsal
part, but length of hairs on middle part of meso-
pleura short, distinctly shorter than that of the
dark lateral band (which is sometimes lacking)
(this character is not valid in a population from
Viet Nam); paramere in distal part strongly
widened and with a distinct tubercle (figs. 102-
113); fore femur slightly flattened, at least basally
(Sri Lanka, India, Thailand, Viet Nam, Malaya,
Borneo, Sumatra, Java) V. modulatus
- Antennal segment 1 usually stouter and thick-
ened in middle of its length, 9.5-11.5 times as
long as maximum width in V. hungerfordi and V.
polhemorum, 10.5-12.5 times as long as maxi-
mum width in V. werneri and V. pilosus. (but
these species out of distribution of V. modulatus);
thorax clothed with dense, long pubescence,
length of hairs on mesopleura long, in its middle
(in yellowish part) nearly of same length than that
of dark lateral band which is always developed;
parameres varied; fore femur not flattened 8
8. Parameres distally strongly widened and with a
distinct tubercle in most specimens (fig. 97),
body depressed (Thailand, Malaya)
V. hungerfordi
- Parameres distally only slightly widened, without
or with a faint indication of a tubercle, body de-
pressed or not 9
9. Paramere in middle part broad, about 0.7 times
as broad as width of apical part, usually without a
dimple at base of apical part (fig. 98), body more
depressed, on average first antennal segment
stouter (Borneo: Sabah, Sarawak)
V. polhemorum
- Paramere in middle part very slender, about half
as broad as width of apical part, in most speci-
mens with a very faint dimple at base of apical
part, body less depressed, first antennal segment
usually less thickened 10
10. Paramere slightly twisted (fig. 100) (Indonesia:
Sumba, Sumbawa) V. pilosus
- Paramere straight, not twisted (fig. 99) (Philip-
pines: Palawan Province) V. werneri
147
Tijdschrift voor Entomologie, volume ui, 1998
Map 1 . Ventidius (s. str.) aquarius-group.
± aquarius
■ usingeri
O harrisoni
• malayensis
•k longitarsus
CX3*£S
ds
* We were not able to check the type material of V. dis-
tanti and V. sushmae. Therefore, they are absent from the
key. But according to the original descriptions and the illus-
trations, they both key out with V. modulatus.
The Ventidius aquarius-group
Diagnosis. - Comparatively larger size, length
3.38-4.60 (<?), 3.20-4.40 (9); width 2.32-3.10 (<J),
2.30-2.96 ( 9 ); with distinctively long and slender an-
tennae; segment 1 shorter or subequal to segments 2
and 3 together, usually more than 1 .20 mm; segment
2 in male usually much longer than segment 3 (except
in V. harrisoni); antennal segment 1 with more than 8
long setae, in males segments 2 and 3 with dense erect
hairs and segment 3 with a more or less distinct in-
dentation near apex, which in some species is sur-
rounded by a number of long thin setae apically; in-
ner surface of fore tibia with dense and long setae, in
distal part with a row of stout spinules forming an S-
shaped comb, and in some species with a long row of
stout spinules nearly all over its length; middle and
hind legs distinctively stout and shining, with scat-
tered long spines on femora and tibiae; segment 8 (s8)
of male with a tuft of dark setae at each posterior an-
gle, parameres small and symmetrical, straight or cu-
cumber-shaped, never dilated apically; in three
species endosoma with a third pair of lateral sclerites,
which are very broad but with their distal parts slen-
der and recurved backwards.
Distribution (map 1). - Widely distributed, from
Sri Lanka to the Philippines and Borneo, not reach-
ing Wallacea. Five species with allopatric distribution
except V. malayensis and V. harrisoni which both in-
habit the Southeast Asian mainland and Borneo.
1. Ventidius (s. str.) aquarius Distant
(figs. 7-9, 12, 18, 23, 24, 29, 30, 39, 43, 48, map 1)
Ventidius aquarius Distant, 1910a: 150 (descr. of genus);
Distant 1910b: 157-158 (descr., illustr. of species).
Hungerford & Matsuda 1960: 324-325, 337 (descr., il-
lustr., key). Thirumalai 1986: 10, 28-30 (list, descr., il-
lustr.).
Type locality. — s. india: Travancore.
Type material examined. - Holotype 9 , apt., 's. india:
Travancore, Pallode, 20 miles N.E. of Trivandrum (Annan-
dale)', 'on road side jungle stream (N. Annandale)' (bmnh).
Other material examined. - india (South): 6c? 15 9 apt.,
2c? 59 macr., Karnataka, Gersoppa, Jog Falls, 600 m, 19.-
24.xi.1977, Zool. Mus. Copenhagen Exped. (zmuc,
nhmw); 2 c? , 2 9 apt., Karnataka, Mudigere area, ca. 900 m,
2.-10.XÌ.1977, Zool. Mus. Copenhagen Exped. (zmuc,
ppcc); 2c? 19 apt., Ammattì (S. Coorg), 3100 ft.,
10.V.1951, leg. P.S. Nathan (jtpc); 1 9 apt., same locality,
11.1952 (jtpc); 2c? 5 9 apt., 2 c? macr. Kerala, 15 km SW
Munnar, Kallar Valley, 1250 m, 1.-9.V.1997, 10°02'N
76°58'E, leg. Dembicky & Pacholatko (nhmw, ppcc). sri
lanka: 2c? apt., Ratnapura, 20.ii.196l, leg. K.L.A. Perera
(semc); 7c? 3 9 apt., 2 larvae, Ambagaspihya, l.ii.1961, leg.
K.L.A. Perera (semc, jtpc); 4c? 1 9 apt., 1 <? macr., Hinidu-
ma, Eia, 29.Ì.1958, leg. K.L.A. Perera (semc); lc? 10 9 apt.,
148
Chen & Zettel: Revision ofVentidius
South Prov., Udugama, N of Galle, 50 m, 22.x. 1974, leg.
N.M. Andersen (zmuc, nhmw); 5c? 2 $ apt., 2$ macr.,
South Prov., Homadola, Udugama, Kan-eliya, from river
pools, leg. P.B. Karunaratne (jtpc, nhmw); 1 $ macr., Uva
Prov., Helagama, Meemina Hela-Large, pool at forest of
hill, 500 ft., 7.-8.XÜ.1967, leg. P.B. Karunaratne (jtpc).
Redescription
Dimensions. - Apterous form, length 3.48 (<?),
3.62 (9), width 2.40 (c3), 2.55 (9), width of head
1.31 (â), 1.31 (9).
Colour (fig. 7). - Whole body prominently pale,
with distinct dark stripes and marks. Eyes mainly
dark brown to black, their inner margin brownish.
Interoculus pale and its dark marks faint, along inner
margin of eye with broad dark stripes reaching the
posterior margin of vertex. Antennal segments dark,
except basal 1/5 of segment 1. Pronotum yellowish,
its anterior margin dark and with dark lateral marks.
Mesonotum with dark lateral stripes. Posterior mar-
gin of metanotum dark and with a small triangular
dark mark basomedially which is confluent with the
dark mark of tergites, and with two more or less
square dark lateral marks which are confluent with
the dark strip of mesonotum; a transverse dark stripe
along anterior margin of acetabula which broadens at
external angle of mesonotum. Mesopleura yellowish
with a broad longitudinal stripe running through its
length, not reaching its anterior margin (fig. 12). An-
terior margin and external angle of metacetabula dark
(fig. 29, 30). Fore leg dark, femur with basal Vi yel-
lowish (fig. 23). Middle and hind leg dark. Tergite 1
with two yellowish marks laterally, tergites 2-3 dark
with a triangle-shaped yellowish mark in the centre,
4-8 yellowish with posterior margin dark. Lateroter-
gites yellowish with dark posterior margins. Connex-
iva dark. Venter yellowish.
Pilosity. - Around antennal segment 1 scattered
12-14 long spinules, and with 7-8 distal short spin-
ules, segment 2 with 6 spinules along its length, seg-
ment 3 with one distal spinule in female and a tuft of
longer setae around the distal part in male, in distal
part of segment 1 and all over the length of segments
2 and 3 ventrally with short erect dark hairs (fig. 18).
Dorsum and pleura bearing dark pubescence, which
is especially prominent on head. About 1 5 long setae
along inner surface of fore femur, their lengths de-
creasing from basal to distal part. Fore tibia clothed
by dense dark setae which form the whole tibia as a
brush, but especially dense in distal part, and there
with an S-shaped comb (fig. 23), in male along inter-
nal face with a row of stout spinules which are in-
creasing in length and becoming more slender to-
wards distal part. Long stiff spines scattered along
middle and hind legs, tibiae with short stiff spinules.
Basal part of middle and hind femora each with two
very long trichobothria-like setae on dorsal surface,
the external one shorter than internal one. The tri-
chobothria-like setae were lost in most specimens we
have seen. Venter clothed by golden pubescence, on
abdominal segments the pilosity longer and denser,
ventral face of segment 8 of male with tufts of broad-
er ranged (comparing with that in V. harrisonî) dark
setae at each posterior angle (fig. 39).
Structural characters. - Interoculus narrower than
width of an eye, 0.45 : 0.42 in males, 0.48 : 0.41 in
females. Antennae distinctly long and slender, espe-
cially the second segment in male much longer than
in most species (measurements see table 1); segment 3
modified at subdistal part, near apex with a deep in-
dentation (fig. 18). Mesonotum swollen, its lateral
width 1.52 (cT), 1.70 (9). Intersegmental suture be-
tween meso- and metanotum distinct. Metanotum
somewhat declivent. Lower half of hind margin of
metacetabula rounded (slightly bilobate in caudal
view), broad and nearly transverse in dorsal view (figs.
29, 30). Metasternal tubercle prominent in male. An-
terior margin of abdominal tergite 1 visible; anterior
margin of tergite 2 faint and obliterated strongly at
medium part; anterior margin of tergite 3 arched for-
wards. Abdominal sternite 7 as long as the preceding
abdominal sternites together in both sexes.
Male terminalia. - Parameres (fig. 43) small, sym-
metrical, more or less straight, cucumber-shaped, not
extending beyond genital segments, with blunt apex.
Endosoma (fig. 48): dorsal sclerite long and recurved
proximally, ventral sclerite long, lateral sclerite
straight, broadened at two ends, second lateral scle-
rites distinct, apical sclerite indistinct.
Female terminalia. - Abdominal sternite 7 large el-
evated medially, posterior margin concave forward.
Macropterous form. - Same as above with the fol-
lowing exceptions: apex of pronotum pointed (mea-
surements see table 3); length of fore wings from
humeri to apex 4. 1 2. Yellowish mark of pronotum see
figure 9.
Distribution (map 1). - S. India: Kerala. Sri Lanka:
Sabaragamuwa (Ratnapura), Southern (Hiniduma),
Prov.? (Ambagaspihya) .
Comparative notes. - According to the relatively
short segment 2 of fore tarsus, the row of stiff spines
on fore tibia of males, and the shorter apical setae on
segment 3 of antenna of males, V. aquarius is closely
related to V. barrisoni and V. usingeri. The other two
species of the group, V. malayensis and V. longitarsus
sp. n., are easily distinguished by the long fore tarsi in
both sexes. Ventidius aquarius shares the deep inden-
tation in male segment 3 of antenna with V. usingeri,
but differences are found in colour, in the deep semi-
circular incision ventrally on segment 8 of males, and
in the shape and absence of pubescence on the para-
meres.
149
Tijdschrift voor Entomologie, volume hi, 1998
Figs. 23-28. Dorsal view of right fore legs. - 23, Ventidius aquarius (male); 24, same species (female); 25, V. usingeri (male);
26, V. harrisoni (male, paratype); 27, V. malayens is (male); 28, V. longitarsus (male, holotype).
2. Ventidius (s. str.) usingeri Hungerford & Matsuda
(figs. 6, 13, 19, 25, 31, 32, 40, 44, 49, map 1)
Ventidius usingeri Hungerford & Matsuda, 1960: 326-327
(descr., illustr., key).
Type locality. - philippines: Los Banos (Luzon).
Type material examined. - philippines: holotype <?,
macr., 'Los Banos, P.i., 17.vii.1936', 'R.L. Usinger collec-
tor', formerly belonged to Usinger collection (cnhm); allo-
type 9, macr., 'cNHM-Philippines Zool., Exped. (1946-
1947)', 'Borungkot, Upi, Cotabato Province, Mindanao
1500 ft. '47', and 'stream through original forest' (cnhm).
Other material examined. - philippines: 1 1 â 7 9 apt., 2â
2 9 macr., Quezon Province, W. Atimonan, Quezon NP, Old
Zigzag Road, 12.ii.1996, leg. H. Zettel (zcwa, gclb, nhmw,
ppcc, uplb); lc? 79 apt., Luzon, Quezon Province, Quezon
N.P., stream at entrance house, CL 1969, lO.vii. 1985, leg. J.T.
& D.A. Polhemus (jtpc, ppcc); 1 S 1 9 apt., 1 3 1 9 macr.,
Luzon, Quezon Province, Quezon N.P., Nalubog Creek, CL
1971, 10.vii.1985, leg. J.T. & DA Polhemus (jtpc); 1 9 apt.,
Polillo Island, leg. Baker (fmhf); 30 49 apt., 1 c? macr., Min-
danao, Bukidnon Prov., Malaybalay, Kaamulan Site, 650 m,
12.11.1996, leg. H. Zettel (90d) (nhmw, uplb, ppcc); 19
macr., Bukidnon Prov., Malaybalay, Kaamulan Site, 650 m,
6.-7.1 1.1996, leg. H. Zettel (90c) (nhmw).
Redescription
Dimensions. - Body length 3.65 (â), 3.79 (9),
width 2.55 (c?), 2.68 (9), width of head 1.40 (â),
1.40(9).
Colour (fig. 6). - Whole body prominently yellow-
ish, dorsal ground colour yellowish with distinct dark
marks. Eye dark brown with grey lines. Interoculus
pale, with a crescent dark mark along inner margin of
eye in male, prominently smokey darkened in female,
and the outline of dark mark rather obscure. Anten-
nal segments black to brownish, except basal % of
segment 1 yellowish. Pronotum and mesonotum yel-
lowish with dark lateral stripes. Metanotum dark at
basal half and with a median triangular dark mark.
Mesopleura with a brownish stripe running through
its length, not reaching its anterior margin but con-
nected with anteroexternal angle of metacetabula (fig.
13). Metacetabula with two yellowish marks (figs. 31,
32). Fore leg with yellowish fore femur, but apical 2/5
more or less dark (fig. 25); tibia and tarsi dark. Mid-
dle and hind femora in male brownish, yellowish
basally. Abdominal tergites and laterotergites in some
specimens prominently yellowish; in some specimens
150
Chen & Zettel: Revision ofVentidius
mainly dark. Connexiva 1-3 dark, the rest yellowish.
Venter yellowish.
Pilosity. - Trichobothria on vertex conspicuous.
Around antennal segment 1 with 10-12 long stiff dark
spines; inner surface of segments 2-4 with dark short
erect hairs (fig. 19). The hair around the constriction
of male antennal segment 3 not specially modified.
Pronotum and fore wing bearing dark pilosity. Pleura
clothed with dark pubescence. Venter clothed by
golden pubescence. Inner surface of male fore femur
with 5-7 stiff spines except the long trichobothria-like
pilosity; fore tibia clothed by dense dark setae which
form the whole tibia as a brush, its inner surface in
males with a dense row of dark spinules which short
and stout at base and gradually becoming longer and
thinner towards apex of tibia. Dark stiff spines scat-
tered along middle and hind legs. Abdominal segment
8 of males with a tuft of dark setae at each corner of
hind margin ventrally (fig. 40).
Structural characters. - Interoculus subequal to
width of an eye, 0.51 : 0.48 in male, 0.47 : 0.46 in fe-
male. Antennae long, segment 2 much longer than 3
in male, slightly longer than 3 in female; subdistal
part of the segment 3 constricted (fig. 1 9) in male
(measurements see table 1). Pronotum more or less
triangular (measurements see table 3). Lateral width
of body 1.45 ((?), 1.66 (9). Lower part of metac-
etabula obtusely triangular in caudal view (figs. 31,
32). Length of wing 3.00 (cT), 3.50 (9). Metaster-
num keel-shaped, distinctly projecting backwards.
Fore femur slender and not modified, claws rising
from Vi of second tarsal segment. Anterior margin of
tergite 1 indistinct, anterior margin of tergite 2 faint
and obliterated strongly at medium part, anterior
margin of tergi tes 3 arched forwards. Abdominal ster-
nite 7 as long as the preceding abdominal sternites to-
gether in both sexes.
Male terminalia. - Parameres (fig. 44) very small,
symmetrical, both bar-shaped, with blunt apex; not
extending beyond genital segments. Endosoma (fig.
49): dorsal sclerite long and recurved proximally, ven-
tral sclerite long, lateral sclerite more or less straight,
broadened and curved at two ends, second lateral
sclerites long and thin, third lateral sclerites irregular-
ly shaped, apical sclerite not visible.
Macropterous form. - Same as above with the fol-
lowing exceptions: apex of pronotum pointed (mea-
surements see table 3); length of fore wings from
humeri to apex 3.00 (S) 3.40 (9). Pronotum yel-
lowish with two dark marks at anterior margin and
one thick transverse band between humeral angles
which is interrupted at median line, lower part of
pronotum with dark margins.
Distribution (map 1). - Philippines: Luzon (Lagu-
na, Quezon), Pollilo Island, Mindanao (Cotabato
South, Bukidnon).
Figs. 29-38. Right metacetabula of female; odd numbers: dorsal view, even numbers: dorsolateral view. — 29, 30, V. aquar-
ius; 31, 32, V. usingeri; 33, 34, V. harrisoni (paratype); 35, 36, V. malayensis; 37 ', 38, V. longitarsus.
151
Tijdschrift voor Entomologie, volume i4i, 1998
40
41
43
44 45
46
Figs. 39-47. - 39-42, Ventral view of male abdominal stemites VIII, showing the pilosity; 43-46. External view of left para-
mere. - 39, 43, Ventidius aquariusr, 40, 44, V. usingeri; 41, 45, V. harrisoni (pararype); 42, 46, V. malayensis, 47, V. longitar-
sus (pararype).
Comparative notes. - This species is very similar to
V. aquarius (from South India and Sri Lanka), for dif-
ferences see under the comparative notes for that
species.
3. Ventidius (s. str.) harrisoni Cheng
(figs. 14, 20, 26, 33, 34, 41, 45, 50, map 1)
Ventidius harrisoni Cheng, 1965: 155-158, 163 (descr., il-
lustr., key); Kovac & Yang 1989: 285 (ree).
Type locality. - Singapore.
Type material examined. - Holotype c?, apt., Singapore:
Seletar River (Sungai Selerar) (1°24'N., 103°47'50' E.),
10.iv.1964 (bmnh). Paratypes, 5<? 59, apt., same locality
data as holotype (bmnh, zrcs).
Other material examined. - Singapore: 1 1 <? 9 9 apt., 46
larvae, Lower Peirce, forest stream, J, Dhm Phy, 6.xi.l991,
coll. CM. Yang (zrcs, ppcc, nhmw); 10c? 15$ apt., Mac
Ritchie, up stream, 17.vi.1994, NSI6IA (zrcs, ppcc,
nhmw); 1 cT 69 apt., 3 larvae, Nee Soon, Swamp forest,
13.vi.1994, NS159B (zrcs); 6<? 11 9 apt., 5 lv.,
MacRitschie Reservoir, SICC nr. plot 4, 28.V.1993, leg.
CM. Yang et al., NS104 (zrcs, nhmw); 10c? 139 apt.,
Nee Soon swamp forest and drain, 13.x. 1986, CL 2214, leg.
J.T. & D.A. Polhemus (jtpc, nhmw); 1 9 apt., Chestnut
Drive, 5.V.1994, NS128A (zrcs); Malaysia: 6c? 79 apt.,
North Selangor, Peat Swamp Forest, stream at 0.2 km from
45 km mark road ro Sungei Bosar., 17.vi.1991, 152 (zrcs,
ppcc); 6c? 49 apt., 1 larva, North Selangor, Peat Swamp,
Foiest stream, at 50 km mark to Tanjong Malim,
18.vi.199L 156, leg. H.K. Lua, Mala (zrcs, nhmw); lc?
2 9 apt., 4 larvae, Selangor, Sabak Bernam, 43 km to sg. Be-
sar fm. Tg. Malim, coll. T.R. Lim (zrcs); 3c? 16 9 apt., 13
larvae, Selangor, peat swamp forest, 25.iii.1994, leg. K.L.
Yeo (zrcs, nhmw); 3 c? 79 apt., Johor, Gunong Panri,
track 270, 22.Ì.199L Y734, Leg. K.L. Yeo (zrcs, nhmw);
1 9 apt., Johor, Sg. Temetang, between Kota Tinggi and
Jemelung, 28.4.1993, Y828, leg. CM: Yang (zrcs); 2c?
apt., Johor, Tg. Sedili, Sg. Selangi, muddy water, sandy bor-
tom, up to 1.5 feet water, Coll. K.L. Yeo & Chia Yi
22.iv.1992 (zrcs); 2c? apt., Johor, swamp forest stream, 61
km NE of Johor Bharu on Mersing road, 1 6.x. 1986, CL
2220, J.T. & D.A. Polhemus (jtpc); lc? 1 9 apt., Johor,
swamp foresr srream, 15 km W of Sedili Besar, 20 m,
16.x. 1986, CL 2218, leg. J.T. & D.A. Polhemus (jtpc); 1 c?
19 apt., Johor, Sg. Mupoh, 14.X.1964, leg. L. Cheng,
ZRCS&.647 1-6472 (zrcs); lc? apt., 788L, Johor, S. Bong,
5.iv.l992, leg. K.L. Yeo (zrcs); lc? 1 9, apt., Terengganu,
north of Ayer Puteh, about 121 km on road from Kuanran
ro Kuala Terengganu, 19.iii.1992, L181, leg. H.K. Lua
(zrcs); 6c?, 39 apt., Kelantan, waterfall 10 km NWofPasir
Puteh, CL 2084, 21.viii.1985, leg. J.T. & D.A. Polhemus
(jtpc, nhmw); 3c? 49, apt., Sarawak, Semengoh, NSG, 30
km S. Kuching, 17.ii.1993, leg. H. Zettel (2) (nhmw,
ppcc); 1 S 39 apt., Sarawak (Borneo), Rumah Kabau anak
152
Chen & Zettel: Revision ofVentidius
muggot, Ng sebong Baleh, 25 km E. Kapit, III. 1994, leg. J.
Kodada (nhmw); 1 â apt., Sarawak, ca. 40 km SE Kapit,
Rumah Ugap Ng, marating bena Kapit, III. 1994, leg. J. Ko-
dada (nhmw); \6 apt., Sarawak (Borneo), Tapah Riv., 16
km NW of Bau, CL 2052, 10.viii.l985, leg. J.T. & D.A.
Polhemus (jtpc); 28 49 apt., Sarawak (Borneo), 4 km S
Tebakang, 19-viii. 1 985, CL 2044, leg. J.T. & D.A. Polhe-
mus (jTpc, ppcc); 1 â apt., Sarawak (Borneo), Kampong
Pueh, Lundu District, 690-1500 m, 6.-12.vi.l958, leg. T.C.
Maa (bimc); 1 9 apt., Sarawak, Kapit District, Merirai V.,
30-300 m, l.-6.viii.l958, secondary forest, leg. T.C. Maa
(bimc); \â apt., Sarawak (Borneo), 8 km S of Tebakang,
stream dissappearing into cave, CL 2046, 9-viii. 1985, leg.
J.T. & D.A. Polhemus (jtpc); Indonesia: 4cv, 69 apt.,
Kalimantan Timur, Long Bagun Ulu, 14. i. 1995, leg. Seyfert
& Graindl (9) (nhmw, ppcc).
Redescription
Dimensions. - apterous form, length 3.38 (â),
3.20 (9), width 2.32 (S), 2.30 (9), width of head
1.30 (c?), 1.26(9).
Colour. - Whole body prominently pale, with dis-
tinct dark stripes and marks. Eyes dark brown, their
inner margin blackish. Interoculus pale, along inner
margin of eye with dark stripe reaching the posterior
margin of vertex. Antennal segments dark, except
basal 1/5 of segment 1. Pronotum yellowish, its ante-
rior margin dark and with lateral dark marks.
Mesonotum with lateral dark stripes. Posterior mar-
gin of metanotum dark and with a triangular dark
mark basomedially which is confluent with the dark
mark of abdominal tergites. Mesopleura yellowish
with a broad longitudinal stripe running through its
length (fig. 14). Dark mark of metacetabula see figs.
33, 34. Fore leg dark, femur with basal 2A yellowish
(fig. 26). Middle and hind leg dark. All tergites and
laterotergites yellowish with posterior margin dark in
female, male tergite 8 totally dark. Connexiva 1-4
dark. Venter yellowish.
Pilosity. - Around antennal segment 1 12-14 scat-
tered long spines and with 7-8 distal short spines, seg-
ment 2 with 3 spines, segment 3 with one distal spin-
ule in female and a tuft of longer setae around the
distal part in male, all antennal segments with dense
dark short erect hairs (fig. 20). Dorsum and pleura
bearing dark pubescence, which is especially promi-
nent on head. More than 10 long setae along inner sur-
face of fore femur, their lengths decrease from basal to
distal part; fore tibia clothed by dense dark setae which
form the whole tibia as a brush, its distal part with an
S-shaped comb (fig. 20), internal face in males with a
dense row of short and stout spinules which are only
slightly increasing in length towards apex. Long stiff
spines scattered along middle and hind legs, tibiae with
short stiff spinules. Venter clothed by golden pubes-
cence, on abdominal sternites the pilosity longer and
denser, ventral face of segment 8 of male with a narrow
tuft of dark setae at each posterior angle (fig. 41).
Structural characters. - Interoculus equal to width
of an eye, 0.44 and 0.44 in males, 0.42 and 0.42 in fe-
males. Antennae distinctly long and slender, especial-
ly the second segment in male much longer than usu-
al (measurements see table 1); segment 3 modified at
distal part, near apex with a shallow indentation (fig.
20). Mesonotum swollen, its lateral width 1.47 (c?),
1.60 (9). Intersegmental suture between meso- and
metanotum distinct and marked by a brown line.
Metanotum somewhat declivent. Lower half of hind
margin of metacetabula slightly bilobate, broad and
nearly transverse from dorsal view (figs. 33, 34).
Metasternal tubercle prominent in male. Anterior
margin of abdominal tergite 1 visible; anterior margin
of tergite 2 clear and obliterated strongly at medium;
anterior margin of tergite 3 arched forwards. Abdom-
inal sternite 7 as long as the preceding abdominal
sternites together in both sexes. Laterotergites broad.
Male terminalia. - Parameres (fig. 45) small, sym-
metrical, slightly hooked, slightly curved upwards,
not extending beyond genital segments, with blunt
apex. Endosoma (fig. 50): dorsal sclerite long and re-
curved proximally, ventral sclerite long, lateral sclerite
straight, broadened at two ends.
Female terminalia. - Abdominal sternite 7 large,
elevated medially, posterior margin concave forward.
Macropterous form. - Unknown.
Distribution (map 1). - Malaysia: Selangor, Johor,
Trenganu, Kelantan, Sarawak; Singapore; Indonesia:
Kalimantan Timur.
Comparative notes. - Ventidius harrisoni is closely
related to V. aquarius and V. usingeri by the characters
mentioned under the comparative notes of V. aquar-
ius. Differences to these species are found in males:
the antennal segment 3 which has only a very shallow
indentation subapically, the spines of the clasping
comb on fore tibia which are only slightly increasing
in length from base to tip and becoming only slightly
thinner towards apex of tibia, and the segment 8 with
a small tuft of hairs laterally. In some samples we
found V. harrisoni mixed with V. malayensis, from
which it differs in length of fore tarsi in both sexes.
4. Ventidius (s.str.) malayensis Hungerford & Matsuda
(figs. 10, 11, 15, 21,27, 35, 36, 42, 46, 51, map 1)
Ventidius malayensis Hungerford & Matsuda, 1960: 325-
326, 336 (descr., illustr., key); Kovac & Yang 1989: 285
(ree).
Type locality. - Malaysia: Selangor.
Type material examined. - Holotype â and allotype 9 ,
Malaya, Selangor, F.M.S., Sungai Ampang, 15.Aug.1926, C.
Dover, Ex. Coll. F.M.S: Museum (bmnh). The holotype also
labelled 'Ventidius aquarius Dist., det. Teiso Esaki', the allo-
type bears the label ' Metrocoris aquarius Dist., det. Dover 26'.
153
Tijdschrift voor Entomologie, volume i4i, 1998
Figs. 48-52. Lateral view of endosoma sclerites. - 48, Ventidius aquarius; 49, V. usingeri; 50, V. harrisoni (paratype); 51, V.
malayensis; 52, V. longi 'tarsus (para type).
Other material examined. -Thailand: 1 <? 1 9 apt., Kan-
chanaburi, Sai Yok waterfall, river Kwai, VIII. 1979, leg. B.
Petersen (zmuc); 4 9 apt., Kanchanaburi, Sai Yok, waterfall,
21.-22.xii.1981, leg. N.M. Andersen (zmuc); Singapore:
4c? 1? apt., 16,5 miles Mersing, Mersing Kluang Rd.,
4.viii.l963, leg. L. Cheng (bmnh); 1? apt., Nea Soon,
swamp forest, 13.vi.1994, NS159B (zrcs); Malaysia: 3c?
7$, lv, Pahang, S. Seladang, 13. and 19.vii.1989, leg. K.L.
Yeo & E. Koh, zrcs 6.2619-2632 (zrcs, nhmw); 2 9 apt.,
Pahang, S. Kernam, 19.vii.1989, leg. K.L. Yeo, zrcs
6.2634-2635 (zrcs); 1? apt., Pahang, S. Kinchin,
13.vi.1989, leg. CM. Yang (zrcs); lc? apt., Johor, Taku,
19.vii.1989, leg. K.L. Yeo, zrcs 6.2633 (zrcs); lc? apt.,
51v, Johor, S. Marong, 5.iv.l992, 788M, leg. K.L. Yeo
(zrcs); lc? 45 apt., 788L, Johor, S. Bong, 5.iv.l992, leg.
K.L. Yeo (zrcs, nhmw); 1 c? 1 $ apt., Johor, stream 12 km
N of Labis, CL 2087, 22.viii.1985, leg. J.T. & D.A. Polhe-
mus (jtpc); 6 c?, 9 9 apt., Johor, 27 km S of Mersing, slow
shaded stream, CL 2058, l4.viii.1985, leg. J.T. & D.A. Pol-
hemus (jtpc, ppcc); 49 apt., Johor, Endau, Sg. Anakjasin,
4.iv.l992, L186, leg. H.K. Lua (zrcs); lc? 39 apt. 5 lv., Jo-
hor, S. Anak, Jasin, Y788D2, leg. K.L. Yeo, 4.iv.l992
(zrcs); 5c?, 5 9 apt, lc? macr., Perak, stream 58 km S of
Grik, CL 2077, 19.viii.1985, leg. J.T. & D.A. Polhemus
(jtpc, nhmw); 2 9 apt., Upper Perak, Longgong, surface of
irrigation channel, fast water, 22.x.-5.xi.l926 (bmnh); 2 9
apt., Sarawak (12), Kelabit Highland, 5 km E Bareo, Pa
Ukat 1000 m, 27.ii.1993 (a) mäandrierender ca 6 m breiter
Fluß, leg. H.Zettel (nhmw); 2 9 apt., Sarawak, Mulu N.P.,
3-5.iii.1993, leg. H. Zettel (14), (e) rechter Zufluß des Tu-
toli River bei Long Iman, ca. 8 m breit, 4.3. (nhmw); 1 c?
1 9 apt., Sarawak, Nanga Pelagus nr. Kapit, 180-585 m, 7-
17.viii.1958, T.C. Maa Collector (bimc); 1 9 apt., Sarawak,
Sadong, Kampong Tapuh 300-450 m, 10.vii.1958, T.C.
Maa, Collector, BISHOP (bimc); lc? 49 apt., Sarawak,
Kapit Dist., Merirai V., 30-300 m, l-6.viii.1958, Secondary
Forest, T.C. Maa Collector, No. MB164 (bimc, semc); 4c?
apt., Sarawak, Sameran River, 2 km W of Tubeh, CL 2047,
19.viii.1985, leg. J.T. & D.A. Polhemus (jtpc); 9c? 109
apt., Sarawak (Borneo), Tapah Riv., 16 km NW of Bau, CL
2052, 10.viii.1985, leg. J.T. & D.A. Polhemus (jtpc,
nhmw, ppcc); 1 9 apt., Sabah, Tibow Estate, slow flow
stream, MB42, 25.5.1996, leg. T.B. Lim (zrcs); Brunei:
1 c? apt., Temburong, Belalong Field Res. Centre, Sungai
Belalong, 60 m, 2-8.V.1995, leg. E. Heiss (zcwa); Indone-
sia: 9 c? 239 apt., 4 c? 79 macr., Kalimantan Timur
Province, Borneo, waterfall and stream, 1 1 km NE of
Samarinda, CL 2091, 27.viii.1985, leg. J.T. & D.A. Polhe-
mus (jtpc, nhmw, ppcc); lc? 1 9 apt., C. Borneo, Sg. Bi-
rang, leg. Mjöberg 1925, Coll. Dr. D. MacGillavry (zmac).
Redescription
Dimensions. - Apterous form, length 4.02 (6),
4.26 (?), width 2.90 (<?), 2.93 (9), width of head
1.60(c)), 1.55(5).
154
Chen & Zettel: Revision ofVentidius
Colour. — Whole body prominently pale, with dis-
tinct dark stripes and marks (fig. 11). Eyes mainly
dark brown to blackish, their margins greyish. Inte-
roculus pale with three obscure long dots, which are
not connected with margin of vertex, along inner
margin of eye dark which is confluent with the dark
mark of pronotum. Antennal segments dark, except
basal 2/5 of segment 1. Pronotum yellowish with lat-
eral dark marks which are more prominent anteriorly.
Mesonotum with lateral dark stripes. Metanotum
with a large triangle-shaped dark mark basomedially
which is confluent with the dark mark of tergites, and
with two irregular lateral dark marks which are con-
fluent with the dark stripes of mesonotum. Mesopleu-
ra yellowish with a broad longitudinal stripe running
through its length (fig. 15); external angle of metac-
etabula dark (fig. 35, 36). Fore leg dark, femur with
basal Vi of fore femur yellowish (fig. 27). Middle and
hind legs dark. Tergite 1 along anterior margin yel-
lowish, the triangle-shaped dark mark confluent with
a dark mark of metanotum; tergites 2-8 dark laterally
with a median yellowish mark which varies individu-
ally. Laterotergites yellowish. Connexiva either black
or 1-4 blackish, 5-7 brownish. Venter yellowish.
Pilosity. - Around antennal segment 1 7-8 scat-
tered long spinules, and with 5-6 distal short spinules,
segment 2 with one subbasal spinule, segment 3 with
a tuft of longer setae around the distal part, all anten-
nal segments with silvery fine hair fringe. Dorsum
and pleura bearing dark pubescence, especially
prominent on head. Circa 20 long setae along inner
surface of fore femur, their length decreasing from
base to apex; fore tibia clothed by dense dark setae
which form the whole tibia as a brush, its distal part
with an S-shaped comb (fig. 27). Long stiff spines
scattered along middle and hind legs, tibia with short-
er stiff spinules. Basal part of middle and hind femo-
ra with two very long trichobothria-like setae on dor-
sal surface respectively, external one shorter than
internal one. Venter clothed by golden pubescence,
on genital segments the pilosity slightly longer and
denser, ventral face of segment 8 of male with a tuft
of broader range (compared with the case in V. har-
risont) dark setae at each posterior angle from ventral
view (fig. 42).
Structural characters. - Interoculus subequal to
width of an eye, 0.55 : 0.50 in males, 0.51 : 0.56 in
females. Antennae distinctly long and slender, espe-
cially segment 2 in male much longer than usually
(measurements see table 1); segment 3 modified at
distal half, near apex with a shallow indentation (fig.
21). Thorax bulbous, mesonotum swollen, its lateral
width 1.93 (<?), 1.96 (9). Intersegmental suture be-
tween meso- and metanotum faint but visible.
Metanotum somewhat declivent. Lower half of hind
margin of metacetabula slightly bilobate, broad and
nearly transverse from dorsal view (figs. 35, 36).
Metasternal tubercle prominent in male. Anterior
margin of abdominal tergite 1 visible; anterior mar-
gins of tergites 2 and 3 faint, obliterated medially and
protruding forward; abdominal sternite 7 as long as
the preceding abdominal sternites together in both
sexes. Laterotergites broad.
Male terminalia. - Parameres (fig. 46) small, sym-
metrical, more or less straight, slightly curved up-
wards, not extending beyond genital segments, with
blunt apex. Endosoma (fig. 51): dorsal sclerite long
and recurved proximally; ventral sclerite long; first
lateral sclerites straight, broadened and curved at two
ends; second lateral sclerites slender and long; third
lateral sclerite broad, distal part slender and recurved
backwards; apical sclerite indistinct.
Female terminalia. - Abdominal sternite 7 large,
elevated medially, sometimes keel-like, posterior mar-
gin concave forward.
Macropterous form. - As apterous form with the
following exceptions: apex of pronotum bluntly
pointed (measurements see table 3); length of fore
wings from humeri to apex 3.30. Dark mark of
pronotum see fig. 10.
Variations. - Specimens from Borneo show small
differences in relative lengths of antennal segments
and colour of pronotum; the Bornean population is
regarded as of infrasubspecific rank.
Distribution (map 1). - Thailand: Kanchanaburi;
Malaysia: Selangor, Pahang, Johor, Perak, Sarawak,
Sabah; Singapore; Brunei; Indonesia: Kalimantan
Timur.
Comparative notes. - This species is very closely re-
lated to V. longitarsus sp. n. It differs from the other
species of the group ( V. aquarius, V. harrisoni, and V.
Usingen) by longer fore tarsi, by lacking a row of stiff
spines on male fore tibia, and by long hairs apically
on segment 3 of males. For differences between V.
malayensis and V. longitarsus sp. n. see comparative
notes ofthat species.
5. Ventidius (s. str.) longitarsus sp. n.
(figs. 16, 17, 22, 28, 37, 38, 47, 52, 53, map 1)
Ventidius (s. str.) sp.: Zettel & Chen 1996: 152, 180 (list,
ree).
Type locality. - viet nam: : Da Lat Province, Mdrak, E
of Ban Me Thuot.
Type material. - Holotype: cT, apt., viet nam: :
[Da Lak Province] Mdrak E. of Ban Me Thuot, 4-
600 m, 8-19.xii.1960, CM. Yoshimoto collector
(bimc). - Paratypes: 1 S 2 9 apt., same locality data as
holotype (bimc, nhmw); 26 , 5 9 apt., Gia Lai-Kon-
tum Prov., 40 km NW Ankhe, Buon Luoi, 14°10'N,
108°30'E, 620-750 m, 28.iii.-12.iv.1995, leg. Pa-
cholatko & Dembicky (nhmw, ppcc).
155
Tijdschrift voor Entomologie, volume i4i, 1998
Fig. 53. Ventidius (s. str.) longitarsus, holotype, apterous male, length 4.60 mm.
Etymology. - This species is named after the long
fore tarsus of male.
Description
Dimensions. - apterous form, length 4.60 (â),
4.40 (9), width 3.10 (<J), 2.96 (9), width of head
1.65 (<?), 1.61 (9).
Colour (fig. 53). - Whole body prominently pale,
with distinct dark stripes and marks. Eyes mainly
dark brown to blackish, their margins greyish. Inte-
roculus pale with three obscure long dots, not con-
nected with margin of vertex, along inner margin of
eye dark posteriorly which is confluent with the dark
mark of pronotum. Antennal segments dark, except
basal 2/5 of segment 1. Pronotum yellowish with
dark lateral mark which is more prominent anterior-
ly. Mesonotum with dark lateral mark. Metanotum
with a triangle-shaped dark mark basomedially which
is confluent with the dark mark of tergites, and with
two irregular-shaped dark lateral marks which are
confluent with the dark strip of mesonotum. Meso-
pleura yellowish with a broad longitudinal stripe run-
ning through its length (figs. 16, 17); external angle
of metacetabula dark (figs. 37, 38). Fore leg dark, fe-
mur with basal Vi yellowish (fig. 28). Middle and
hind legs dark. Tergite 1 along anterior margin yel-
lowish, the triangle-shaped dark mark confluent with
dark mark of metanotum; tergites 2-8 dark laterally
156
Chen & Zettel: Revision ofVentidius
with a median yellowish mark which varies individu-
ally. Laterotergites yellowish. Connexiva either total-
ly black or 1-4 brownish and 5-7 yellowish. Venter
yellowish.
Pilosity. - Around antennal segment 1 7-8 scat-
tered long spinules, and with 5-6 distal short spinules,
segment 2 with one subbasal and one subdistal spin-
ules, segment 3 with a tuft of longer setae around the
distal part, all antennal segments with silvery fine hair
fringe (fig. 22). Dorsum and pleura bearing dark pu-
bescence, which is especially prominent on head. Cir-
ca 20 long setae along inner surface of fore femur,
their lengths decrease from basal to distal part; fore
tibia clothed by dense dark setae which form the
whole tibia as a brush, its distal part with an S-shaped
comb (fig. 22). Long stiff spines scattered along mid-
dle and hind legs, tibia with shorter stiff spinules.
Basal part of middle and hind femora with two very
long trichobothria-like setae on dorsal surface respec-
tively, the external one shorter than internal one.
Venter clothed by golden pubescence, on genital seg-
ments the pilosity slightly longer and denser, ventral
face of segment 8 of male with a tuft of broader
ranged (compared with the case in V. harrisoni) dark
setae at each posterior angle from ventral view.
Structural characters. - Interoculus broader than
width of an eye, 0.63 and 0.52 in males, 0.58 and
0.51 in females respectively. Antennae distinctly long
and slender, especially segment 2 in male much
longer than usual (measurements see table 1); seg-
ment 3 modified at distal half, near apex with an in-
dentation (fig. 22). Thorax bulbous, mesonotum
swollen, its lateral width 1.95 (6), 2.00 (9). Inter-
segmental suture between meso- and metanotum
faint but visible. Metanotum somewhat declivent.
Lower half of hind margin of metacetabula slightly
bilobate, broad and nearly transverse from dorsal view
(figs. 37, 38). Metasternal tubercle prominent in
male. Anterior margin of abdominal tergite 1 visible;
anterior margins of tergites 2 and 3 faint, obliterated
medially and protruding forward; abdominal sternite
7 as long as the preceding abdominal sternites togeth-
er in both sexes. Lateral tergite plate broad.
Male terminalia. — Parameres (fig. 47) small, sym-
metrical, more or less straight, inner surface with in-
dentation at basal half, not extending beyond genital
segments, with blunt apex. Endosoma (fig. 52): dor-
sal sclerite long and recurved proximally, ventral scle-
rite long, lateral sclerite straight, broadened at proxi-
mal end, second lateral sclerite long oblique and
distinct, the third lateral sclerite broad, darkened and
curved at distal part, apical sclerite indistinct.
Female terminalia. — Abdominal sternite 7 large,
elevated medially, posterior margin concave forward.
Macropterous form. - Unknown.
Distribution (map 1). - Viet Nam: Da Lak, Già
Lai-Kontum.
Comparative notes. - This species is very similar to
V. malayensis Distant, but has larger size and darker
colouration. Main difference is the very long segment
2 of male fore tarsus (see table 2), with claws arising
from basal 1/6 of the segment.
The Ventidius modulatus-group
Diagnosis. - Comparatively smaller size, length
2.41-3.00 (<?), 2.50-3.10 (9), width 1.64-1.90 (<J),
1.74-2.20 (9); length of antennae intermediate; seg-
ment 1 longer or subequal to segments 2 and 3 to-
gether, usually less than 1.20 mm; length of segment
2 in male not or slightly longer than segment 3 (up to
1.4 times); antennal segment 1 usually with less than
8 long setae, in males segments 2-4 with dense erect
silvery pubescence and segment 3 without indenta-
tion near apex; inner surface of fore tibia with dense
setae, in distal part with a row of stout spinules form-
ing an S-shaped comb, never with a long row of stout
spinules; middle and hind legs distinctly more slender
than in the V. aquarius-group; segment 8 (s8) of male
without tufts of dark setae at each posterior angle,
parameres small and symmetrical, usually distinctly
dilated apically (except in V. henryi); in three species
endosoma with a third pair of lateral sclerites, which
are very broad only at distal part slender and recurved
backwards.
Distribution (maps 2, 3). - Widely distributed in
Oriental region, from Sri Lanka to Philippines (Pala-
wan region) and Borneo, in the east reaching the Wal-
lacea (Sumbawa and Sumba), but unknown from Su-
lawesi.
6. Ventidius (s. str.) henryi Esaki
(figs. 54, 55, 60, 67-69, 75, 76, 88, 96, 1 14, map 2)
Ventidius henryi Esaki, 1928: 509-511 (descr., illustr.);
Lundblad 1933: 372 (list); Hungerford & Matsuda
1960: 327-329 (descr., illustr., key).
Type locality. - sri lanka: Sabaragabuwa.
Type material examined. - Holotype, S apt., sri lanka:
Sabaragabuwa Prov., Kitulgala, 12.iv. 1927,. leg. G.M. Hen-
ry (bmnh). Paratypes, 3? apt., same locality data as holo-
type (bmnh).
Other material examined. - sri lanka: 3 cT 2? apt.,
Sabaragabuwa Prov., Kitulgayala, Dec., 1934 (semc); AS
3$ apt., Sabaragabuwa Prov., 57th mi. Kitulgayala,
23.vii.1966, leg. Karunaratne (jtpc); 3d 3 9 Central Prov.,
6.-7.viii.l966, leg. P.B. Karunaratne (jtpc, nhmw); 2d 3?
apt., 1 larva, Central Prov., Noari Estate, Noari, 10.vi.1966,
leg. D.N. Bartholomaeusz (jtpc, ppcc); 76 69 apt., 19
macr., Southern Prov., Homadola, Udugama, 2.vi.l966,
leg. Karunaratne (semc, nhmw, jtpc); (Prov.?): 2d 19
apt., Pitamba, Eia., 26.Ì.1958, leg. K.L.A. Perera (semc).
157
Tijdschrift voor Entomologie, volume hi, 1998
54
55
Figs. 54-59.
54, Ventidius henryi, holo-
type, dorsal view of apterous
male (legs removed), length
2.44 mm. — 55, Ventidius
henryi dorsal view of macro-
pterous female (appendages
removed). - 56, 57, Macro-
pterous pronotum. - 56, V.
polhemorum; 57, V. werneri.
—58, V. hungerfordi, dorsal
view of right fore wing (after
Lansbury 1990). - 59, V.
modulatus (female), ventral
view of abdominal segments.
58
Redescription
Dimensions. - Apterous form, length 2.46 (â),
2.70 (9), width 1.64 (c?), 1.84 (9), width of head
1.04 (cT), 1.08(9).
Colour (fig. 54). - Whole body predominantly
dark dorsally, with yellowish marks. Eyes blackish.
Interoculus dark from dorsal view, with a thick M-
shaped yellowish mark at its posterior margin, along
inner margin of eye with a dark stripe which is con-
nected with anterior margin of pronotum. Male an-
tennal segments blackish except at basal lA of segment
1 . Pronotum blackish, posterolateral angle with a very
small and indistinct yellowish mark. Mesonotum
with broad dark lateral stripes, which are confluent
with the lateral black stripe of metanotum. Metan-
otum dark laterally and a triangular dark mark baso-
medially which reaches its anterior margin. Meso-
pleura yellowish with a longitudinal stripe at anterior
half (figs. 67-69); posterior margin of metapleura
dark; metacetabula black with central yellowish mark
(figs. 75, 76). Fore leg totally black (fig. 88). Middle
and hind legs dark. Tergites 1-3 blackish, 4 blackish
with a small obscure yellowish mark in the middle, 5-
6 yellowish, 7-8 dark. Laterotergites and connexiva 1 -
4 blackish, 5-7 yellowish in male, predominantly yel-
lowish with dark margins in female. Venter yellowish,
male segment 8 ventrally darker.
Pilosity. - Inner surface of antennal segment 1 with
4 subbasal, and 1 subdistal long spines, along ventral
surface of segments 2-4 clothed with usual pubes-
158
Chen & Zettel: Revision ofVentidius
cence and longer setae; from distal half of segment 1,
through all segments with dark fine hair fringe. Dor-
sum and pleura bearing dark and golden pubescence.
Long stiff bristles scattered along middle and hind
legs, tibiae with shorter stiff bristles. Venter clothed
by golden pubescence, on genital segments the pilos-
ity slightly longer and denser.
Structural characters. - Interoculus broader than
width of an eye, 0.42 : 0.34 in males, 0.45 : 0.36 in
females. Antennae moderately strong in male, seg-
ment 2 roughly of same length as segment 3 in male,
shorter than segment 3 in females (measurements see
table 1) (fig. 60). Pronotum not bulbous, mesonotum
swollen dorsally, its lateral width 1.08 (a), 1.30 (9).
Intersegmental suture between meso- and metan-
otum faint to obscure. Metanotum somewhat de-
clivent. Lower half of hind margin of metacetabula
obtusely triangular (figs. 75, 76). Metasternal tuber-
cle not prominent in male. Anterior margin of ab-
dominal tergite 1 obscure; anterior margins of tergites
2 and 3 faint, obliterated medially and protruding
forward; abdominal sternite 7 as long as the preceding
abdominal sternites together in both sexes.
Male terminalia. - Parameres (fig. 96) symmetrical,
straight, bar-shaped with blunt apex; slightly extend-
ing beyond genital segments. Endosoma (fig. 1 14):
dorsal sclerite long and recurved proximally, ventral
sclerites long, lateral sclerites straight, hooked at apical
end and broadened at distal end, second lateral scle-
rites long and thin, third lateral sclerites curved.
Female terminalia. - Abdominal sternite 7 large,
broadly elevated medially, posterior margin slightly
concave forward.
Macropterous form. - As apterous form except the
apex of pronotum pointed (measurements see table 3);
length of fore wings from humeri to apex 2.00. colour
pattern of pronotum (fig. 55) black with two small
roundish yellowish blotches. Wings anteriorly black
with fine pubescence and with dense longer black hairs.
Posterior membranous part of wings dark brown.
Distribution (map 2). - Sri Lanka: Sabaragabuwa,
Central Prov., Southern Prov., Pitamba (Prov.?).
Comparative notes. - Ventidius henryi differs from
all other species of the V. modulatus-group by much
darker colouration (e.g. the black pronotum and
completely black fore leg) and very different shape of
paramere which is bar-shaped and not broadened in
distal part.
7. Ventidius (s. str.) hungerfordi Cheng
(figs. 58, 61, 62, 77-80, 89, 90, 97, 115, 123, 124,
map 2)
Ventidius hungerfordi Cheng, 1965:158-159, 163 (descr., il-
lustr. & key); Kovac & Yang 1989: 285 (ree).
Type locality. - Malaysia: Johor.
Figs. 60-66. Dorsal view of male right antennae. - 60, Ven-
tidius henryi; 61, V. hungerfordi, apterous; 62, same species,
macropterous; 63, V. polhemorum; 64, V. werneri; 65, V. pi-
losus; 66, V. modulatus.
Ventidius wallacei Lansbury, 1988: 61-65 (synonym, descr.,
illustr.). Syn. n.
Type locality. - Malaysia: Selangor, Johor.
Type material examined. — Ventidius hungerfordi: Holo-
type S , apt., Malaysia: Selangor, Ampang, Sungai Kongsi
Lapan (3°10' N., 101°46'30'E.), 2000 ft., 13.iii.1964, leg.
L. Cheng (bmnh); Pararypes: 1 S apt., same locality data,
ZRCS 6.998 (zrcs); Id apt., Johor, stream at 48th mile,
Mersing-Kluang Road, leg. C.H. Fernando, zrcs 6.997
(zrcs); 1 9 apt., Johor, Sungai Kayu Ara, 5.x. 1964, leg. L.
Cheng (bmnh).
Ventidius wallacei: Holorype d, macr., Malaysia: Mt.
Ophir [= Gunong Ledang, Johor; Yang, pers. comm.] (A.R.
Wallace) Westwood Coll. Oxford (OUMC).
Other material examined. —Thailand: 2d 4 9 apt., Kan-
chanaburi, Khao Phang waterfall, 70 km NW Kanchanaburi,
10.ix.1991, leg. P. Nielsen (zmuc, nhmw); 19 apt., Kan-
chanaburi, Sai Yok, waterfall, 21-22.xii.1981, leg. N.M. An-
dersen (zmuc); w. Malaysia: 7d 69 apt., 1 9 macr., Johor,
Panti, stream, 20.ix.1990, leg. CM. Yang & H.K. Lua, Y29b
(zrcs, nhmw); 3d 79 apt., Johor, same locality data, Y29e
159
Tijdschrift voor Entomologie, volume hi, 1998
%J74
Figs. 67-74. Lateral view of apterous male, showing the colour pattern of pleura. - 67, 68, Ventidius henryi, apterous; 69, same
species, macropterous; 70, V. polhemorum; 7 '1 , V. werneri;72, V. pilosus; 73-74, V. modulates (iectotype) .
75
76
Figs. 75-87. Right metacetabula of male; odd numbers (except 87): dorsal view, even numbers and 87: dorsolateral view. -
75, 76, Ventidius henryi; 77, 78, V. hungerfordi; 79, 80, same species, macropterous; 81, 82, V. polhemorum; 83, 84, V.
werneri; 85, 86, V. pilosus; 87, V. modulatus (lectotype) .
160
Chen & Zettel: Revision ofVentidius
Figs. 88-95. Dorsal view of male right fore legs. - 88, Ventidius henryi; 89, V. hungerfordi; 90, same species, macropterous;
91, V. polhemorum; 92, V. werneri; 93, V. pilosus (male); 94, same species, female; 95, V. modulatus (\ectovype) .
96
Figs. 96-100. External view of left paramere. - 96, Ventidius henryi; 97, V. hungerfordi; 98, V. polhemorum; 99, V. werneri;
100, V. pilosus; 101, V. pilosus, laterointernal view of right paramere.
161
Tijdschrift voor Entomologie, volume 141, 1998
102
103
104
105
106 107
108
109
110
111
112
113
Figs. 102-1 13. - Ventidius modulatus, variation in parameres: external view of left paramere and dorsal view of right paramere
(106). — 102, lectotype (Java); 103, holotype of V. chinai (Selangor); 104, paratype of V. pubescens (Johor); 105, from Suma-
tra; 106, from Sumatra; 107-113, from Thailand: 107, Chiang Mai (Doi Inthanon); 108, Ubon; 109, Chiang Rai (Fang);
110, Kanchanaburi; 111, Khon Kaen; 112, Chiang Mai (Hui Hong Krei); 113, Petchabun.
(zrcs, ppcc); 2c? 4 9 , Johor, Kota Tinggi waterfall, 16 km N J.T. & D.A. Polhemus (jTpc, nhmw); 1 9 apt., Gua Sungei,
of Kota Tinggi, 60 m, 15.X.1986, CL 2217, leg. J.T. & DA
Polhemus (jtpc, nhmw); 1 9 apt., Johor, Kota Tinggi, wa-
terfall, 25.ix.1993, Y838, leg. K.L. Yeo (zrcs); lc? 1 9 apt.,
Johor, S. Bong, 5.iv.l992, Y788L, leg. K.L. Yeo (zrcs); 1 S
1 9 apt., Johor, S. Anak, Jasin, 4.iv.l992, Y788, leg. K.L.
Yeo (zrcs); 1 S macr., Perak, Inah Hill, stream nr. reservoir,
ll.iii.1927, coll. Dover (bmnh); 10 9 apt., Kelantan, water-
fall 10 km NW of Pasir Puteh, CL 2084, 21.viii.1985, leg.
Baling Kedah, ex megaderma spasma medium Gerrid
(6324), 2.V.1935 (FOG23280) (bimc).
Redescription
Dimensions. - Body length 2.77 (<?), 2.60 (9),
width 1.75 (c?), 1.80 (9), width of head 1.01 (<J),
1.10(9).
162
Chen & Zettel: Revision ofVentidius
Figs. 114-122. Lateral view of endosoma sclerites. — 114, Ventidius henryi; 115, V. hungerfordi; 116, V. polhemorum; 117, K
wemeri; 118, V. pilosusr, 119-122. K modulatm. 119, lectotype (Java); 120, paratype of V. pubescenr, 121, From Thailand (Doi
Inthanon); 122, holotype of V. chinai.
Colour (fig. 123). - Whole body prominently yel-
lowish, dorsal ground colour yellowish with distinct
dark marks. Eye dark brown. Interoculus pale, with-
out distinct dark mark. Dark mark along inner mar-
gin of eye broad and reaching posterior margin of ver-
tex. Antennal segments dark, except basal Va of
segment 1 . Pronotum blackish with a triangular yel-
lowish mark medioposteriorly. Mesonotum yellowish
dorsally, with dark lateral stripes. Metanotum yellow-
ish with a small triangular black mark medioposteri-
orly. Mesopleura with a broad brownish stripe run-
ning through its length, lower part of metacetabula
dark (figs. 77, 78). Fore leg dark with basal Vi yellow-
ish (figs. 89, 90). Middle and hind legs dark. Tergites
1-2 completely black, 3 prominently dark with a
small faint yellowish spot in the middle, 4-6 yellowish
but black laterally, 7 yellowish with dark posterior
margin, posterior half of tergite 8 black. Lateroter-
gites 1 -2 completely black, 3 dark with small yellow-
ish spot, 4-5 yellowish with dark margin, 6 yellowish.
Connexiva 1-3 dark, the rest yellowish. Venter yel-
lowish. Metasternum tubercle yellowish.
Pilosity. - Trichobothria on vertex conspicuous.
Inner surface of antennal segment 1 with 4 subbasal,
and 1 subapical long spines; segments 2-4 with scat-
tered dark short setae (fig. 61). Dorsum and pleura
bearing longer dark setae. Pleura covered with black-
ish depressed longer silvery hairs. Venter clothed by
golden pubescence. Stiff spines scattered along mid-
dle and hind legs.
Structural characters. - Interoculus longer than
width of an eye, (0.41 : 0.36) in male, equal (0.40 :
0.40) in female. Antennae not modified, segment 2
longer than 3 in males (fig. 61), roughly of same
length in females (measurements see table 1). Thorax
not bulbous, mesonotum somewhat swollen, its later-
al width 1.10 in both sexes. Intersegmental suture be-
tween meso- and metanotum weak but visible.
Metanotum declivent. Lower part of metacetabulum
obtusely triangular, but bilobate, the internal lobe
smaller than external lobe and curved towards its
body, therefore easily ignored (figs. 77, 78). Fore fe-
mur slender and not modified, claw rising from Vs of
segment 2 of fore tarsus. Anterior margin of abdomi-
nal tergite 1 distinct; anterior margins of tergites 2
and 3 obliterated medially and protruding forward;
abdominal sternite 7 as long as the preceding abdom-
inal sternites together in both sexes.
163
Tijdschrift voor Entomologie, volume mi, 1998
Fig. 123. Ventidius (s. str.) hungerfordi, holotype, apterous
male, length 2.77 mm.
Male terminalia. - Parameres (fìg. 97) symmetrical,
slender and twisted in middle part, distinctly widened
in apical part, usually with distinct small tubercle on
external side of ventral margin, with rounded apex, ex-
tending beyond genital segments; Endosoma (fig.
115): dorsal sclerite long and recurved proximally,
ventral sclerites long, lateral sclerites straight, proximal
parts broadened and hooked, second lateral sclerites
thin, apical sclerite indistinct.
Macropterous form. - We present a redescription
of the macropterous holotype of V. wallacei:
Dimensions. - Body length 2.70, including wing
3.00, width 1.45, width of head 1.09.
Colour (fig. 124). - Whole body prominently yel-
lowish, dorsal ground colour yellowish brown with
distinct dark marks. Eye dark brown. Interoculus
pale, without distinct dark mark. Dark mark along
inner margin of eye reaching posterior margin of ver-
tex. Antennal segments dark, except basal 1/5 of seg-
ment 1 . Pronotum with blackish margins and with a
large central irregularly shaped yellowish spot.
Metanotum dark. Mesopleura yellowish, external an-
gle of metacetabula dark. Fore leg dark with basal Vi
yellowish. Middle and hind legs dark. Wings dam-
aged, remnants dark rich brown, venation slightly
darker, remnants of membranous part not differently
coloured from fore wings. Tergite 1 completely yel-
lowish, tergite 2 dark, tergites 3-7 yellowish, posteri-
or half of tergite 8 dark. Laterotergites yellowish.
Connexiva 1-4 dark, 6-7 yellowish. Venter yellowish.
Metasternum tubercle brownish.
Pilosity. - Inner surface of antennal segment 1 with
3 subbasal, and 1 subapical long spines; segments 2-4
with scattered dark short setae. Dorsum and pleura
bearing dark pubescence. Pleura covered with black-
ish depressed hairs. Venter clothed by golden pubes-
cence, especially on genital segments with longer pi-
losity. Stiff spines scattered along middle and hind
legs. Basal part of middle and hind femur with two
very long trichobothria-like setae on dorsal surface.
Structural characters. - Interoculus broader than
width of an eye, 0.40 and 0.32 respectively. Antennae
not modified, second segment longer than the third
in males (fig. 62), measurements see table 1. Thorax
not bulbous, lateral width of mesonotum 1.10.
Mesosternal ridge almost reaching an imaginary line
through faint brown spots anterior of lateral
mesosternal ridges. Distal ridge slightly raised; lower
part of metacetabula obtusely triangular (figs. 79, 80).
Metasternal tubercle slightly prominent in male. Fore
femur slender and not modified, claw rising from Vi
of segment 2 of fore tarsus. Wing venation see figure
58. Anterior margin of abdominal tergite 1 distinct;
anterior margins of tergites 2 and 3 obliterated medi-
ally and protruding forward; abdominal sternite 7 as
long as the preceding abdominal sternites together in
both sexes. Lateral tergite plate broad.
Male terminalia. - Parameres symmetrical, both
pulpboard-shaped, with blunt apex; both extending
beyond genital segments, internal side near lower
margin with a very small but visible tooth-like tuber-
cle, very easily ignored. Endosoma: dorsal sclerite
long and recurved proximally, ventral sclerites long,
lateral sclerites straight, proximal parts broadened
and hooked, second lateral sclerites thin, apical scle-
rite indistinct.
Distribution (map 2). - Thailand: Kanchanaburi;
Malaysia: Selangor, Johor, Kelantan, Perak.
Remarks. — Ventidius wallacei was described after a
single macropterous male. The dense long pilosity of
mesopleura and the thick antennal segment 1 remind
immediately of V. hungerfordi. Lansbury (1988) com-
pared V. wallacei with V. hungerfordi and states dif-
ferences in colouration and parameres ('dark brown-
black pigmentation of wallacei is much more
extensive than that of hungerfordi; the paramere of the
latter has a small tubercle on the rear margin, wallacei
has the margins evenly rounded'). We cannot follow
this opinion: The dark colouration corresponds well
with a macropterous male of V. hungerfordi from Per-
ak. After dissection of the paramere of the holotype of
V. wallacei, we could detect a faint indication of a tu-
164
Chen & Zettel: Revision ofVentidius
Fig. 124. Ventidius (s. str.)
hungerfordi, macropterous
male, length 3.0 mm (= V.
wallacei Lansbury).
bercle on paramere from a certain direction of view;
the tubercle is also weakly developed in V. hungerfor-
di (compared with V. modulatus). Although the para-
mere of V. wallacei is slightly more slender than is
usual in V. hungerfordi, we regard this character as an
individual variation and subsequently V. wallacei as a
synonym of V. hungerfordi.
Comparative notes. - Ventidius hungerfordi seems
to be closely related to the following three species ( V.
polhemorum sp. n., V. werneri, V. pilosus sp. n.) on the
basis of the long pilosity on mesopleura and by a
more or less enlarged antennal segment 1 in males
(compared with that in V. modulatus). It can be sepa-
rated from all of these species by the more distinct tu-
bercle on the ventral external face of the paramere,
which is lacking or very reduced in the other three
species. Ventidius hungerfordi can be separated from
V. modulatus (which is the only sympatric species of
this group) also by a flatter body in both sexes.
8. Ventidius (s. str.) polhemorum sp. n.
(figs. 56, 63, 70, 81, 82, 91, 98, 116, 125, map 2)
Type locality. - Malaysia: Sabah.
Type material. — Holotype 6 , apt., allotype 9 , apt.,
Malaysia: Sabah, Borneo, 34 km NE of Kota Belud,
CL 2033, 5. VIII. 1985, leg. J.T. & DA Polhemus
(jtpc). - Paratypes: 9c? 26 9 apt., 26 4 ? macr., same
locality data as holotype (jtpc, nhmw, ppcc); 24 c?
52 9 apt., 26 3 9 macr., Sabah, Borneo, 40 km NE of
Kota Belud, CL 2032, 5.viii.l985, leg. J.T. & D.A.
Polhemus (jtpc, nhmw, ppcc); 36 1 9 apt., Sabah,
Borneo, stream 5 km S of Poring Hot Springs, CL
2024, 2.viii.l985, leg. J.T. & DA. Polhemus (jtpc);
1 6 apt., Sabah, (North Borneo, SE), Forest Camp, 19
km N of Kalabakan, 12.xi.1962, light trap [?], leg. Y.
Hiyashima (bimc); 1 9 macr., same locality data,
13.xi.1962 (bimc); 49 apt., Sabah, Danum Valley,
Sapat Kalisan, 12.2.1997, leg. H. Zettel (15) (umsm,
nhmw); 1 6 3 9 apt., Sabah, Danum Valley, Palum
Tambun, 12.2.1997, leg. Zettel et al. (P83 & P90)
(umsm, nhmw, spcm); 36 3 9 apt., 19 macr.,
Sarawak (Borneo), 8 km S of Tebakang, stream disap-
pearing into cave, CL 2046, 9.viii.l985, leg. J.T. &
DA. Polhemus (jtpc, nhmw); 1 6 apt. Sarawak,
Sameran River, 2 km W of Tubeh, CL 2047,
19.viii.1985, leg. J.T. & DA. Polhemus (jtpc); 16
2 9 apt., 19 macr., Sarawak (Borneo), 4 km S
Tebakang, 19.viii.1985, CL 2044, leg. J.T. & D.A.
Polhemus (jtpc).
Etymology. - The species is named after Dr. J.T.
Polhemus (Englewood) and Dr. D.A. Polhemus
(Washington), for their distinguished achievements in
the aquatic and semiaquatic Hemiptera fauna of the
world, and for their generous offering of material de-
posited in the jtpc and bimc collections for our study.
165
Tijdschrift voor Entomologie, volume i4i, 1998
Fig. 125. Ventidius (s. str.) polhemorum, paratype, apterous
male, length 2.78 mm.
Description
Dimensions. - Body length 2.78 (<î), 2.89 (9),
width 1.73 (S), 1.95 (9), width of head 1.10 (cî),
1.10(9).
Colout (fig. 125). - Whole body prominently yel-
lowish, dorsal ground colour yellowish with distinct
dark marks. Eye dark brown. Interoculus pale, central
part with darker pilosity. Dark mark along inner mar-
gin of eye broad and reaching posterior margin of ver-
tex. Antennal segments dark, except basal Va of seg-
ment 1. Pronotum yellowish with dark lateral mark.
Mesonotum yellowish dorsally, with dark lateral
stripe. Metanotum yellowish with a small triangular
black mark medioposteriorly and dark lateral stripes.
Mesopleura yellowish, lower part and external margin
of metacetabula dark (fig. 70). Fore leg dark with basal
half yellowish (fig. 91). Middle and hind legs dark.
Tergi tes 1-2 completely black, 3 prominently dark
with a small faint yellowish spot in the middle, 4 black
with a big yellowish mark in the middle, 5-6 yellowish
but black laterally, tergite 7 yellowish with dark poste-
rior margin, posterior half of tergite 8 black. Lat-
erotergites 1-2 completely black, 3-7 yellowish. Con-
nexiva slightly darkened. Venter yellowish.
Metasternum tubercle yellowish.
Pilosity. - As a whole, body with long, more promi-
nent pilosity. Inner surface of antennal segment 1 with
4 subbasal and 1 subapical spine, and 2-3 long spines
scattered; segments 2-4 with scattered dark short setae,
and especially inner surface of segments 2-4 with fine
hair fringe (fig. 63). Dorsum and pleura bearing dark
setae, more conspicuous on pleura and metacetabula.
Venter clothed by pale pubescence. Genital segments
with longer and darker setae. Stiff spines scattered
along middle and hind legs.
Structural characters. - Interoculus broader than
width of an eye (0.46 : 0.39) in male, equal (0.40 :
0.40) in female. Antennae not modified, segment 2
longer than 3 in males (fig. 63), shorter than in fe-
males (measurements see table 1). Thorax not bul-
bous, mesonotum somewhat swollen, its lateral width
1.20 (â), 1.23 (9). Intersegmental suture between
meso- and metanotum weak but visible, and marked
with a fine brownish line laterally. Metanotum de-
clivent. Lower part of metacetabula obtusely triangu-
lar, but bilobate in caudal view, the internal lobe
much smaller than external lobe and slightly curved
mediad (figs. 81, 82). Metasternal tubercle darkened.
Fore femur slender and not modified, claw rising
from Vi of segment 2 of fore tarsus. Anterior margin
of abdominal tergite 1 faint; anterior margins of ter-
gites 2 and 3 obliterated medially and protruding for-
ward; abdominal sternite 7 as long as the preceding
abdominal sternites together in both sexes.
Male terminalia. - Parameres (fig. 98) symmetri-
cal, curved, broad in middle part, slightly widened
apically, usually without any tubercle, in some speci-
mens with a faint indication of a dimple, with round-
ed apex; extending beyond genital segments. Endoso-
ma (fig. 1 1 6) : dorsal sclerite long and recurved
proximally, ventral sclerites long, lateral sclerite
straight, broadened and hooked at two ends, second
lateral sclerite thin, apical sclerite indistinct.
Macropterous form. - 1.40 mm long, maximum
width 1.51 mm. colour pattern (fig. 56) of pronotum
yellowish anteriorly and in the central area with two
yellowish blotches, lateral and posterior margin black-
ish. Wings anteriorly blackish with fine pubescence
and with scattered longer black hairs. Posterior mem-
branous part of wings dark brown. The other mor-
phological characters as apterous form except the apex
of pronotum pointed (measurements see table 3);
length of fore wings from humeri to apex 2.40 mm.
Distribution (map 2). - East Malaysia: Sabah,
Sarawak.
Comparative notes. - This species is very similar to
V. hungerfordi (Malaya), V. werneri (Philippines:
Palawan), and V. pilosus sp. n. (Indonesia: Sumba) by
colour pattern, conspicuous pilosity, and slightly bilo-
bate lower structure of metacetabula. Ventidius
werneri and V. pilosus sp. n. differ in males by a dis-
tinctly more slender antennal segment 1 and by differ-
166
Chen & Zettel: Revision ofVentidius
Map 2. Ventidius (s. str.) modula-
fttf-group, partim.
• henry i
Q bungerfordi
■ polhemorum
± werneri
•k pilosus
izzz5s*0às
ent shapes of parameres. Ventidius hungerfordi, which
is similar in most characters can be distinguished by
the paramere which in V hungerfordi is strongly
widened apically and usually with a distinct tubercle
on the ventral external face, but in V. polhemorum sp.
n. is less widened (because of the wide middle part)
and usually without any tubercle.
9. Ventidius (s. str.) werneri Hungerford & Matsuda
(figs. 57, 64, 71, 83, 84, 92, 99, 1 17, 126, map 2)
Ventidius werneri Hungerford & Matsuda 1960: 330-331
(descr., illustr., key).
Type locality. - philippines: Palawan.
Type material examined. - Holotype c? apt., and allotype
9 apt., philippines: Palawan 'cnhm Philippines Zool. Ex-
ped. (1946-47). F.G. Werner, Puerto Princesa, Palawan Is.
Sea Level, March (1947)' (cnhm).
Other material examined. - 5<? 15 9, all apt., philip-
pines: Palawan, Sabang, 0-30 m, 27.iii.1994, leg. Zettel
(52b) (nhmw, ppcc); lc? 11 9 apt., Id" macr., Palawan, 9
km W Puerto, Princese, lwahig, Balsahan riv., leg. Zettel,
24.iii.1994 (48, 60) (nhmw, ppcc); 2 9 apt., Palawan, Ba-
cungan Creek, 18 km NW of Puerto Princesa, CL 2019,
28.vii.1985, leg. J.T. & D.A. Polhemus (jtpc); 12c? 169
apt., Palawan, Apoapo Arawan River, 76 km S of Puerto
Princesa, CL 2005, 25.vii.1985, leg. J.T. & D.A. Polhemus
(jtpc, nhmw, ppcc); 6c? 1 1 9 apt., Palawan, Taritien River,
at Trident Mine, 7 km NW of Narra, CL 20 1 1 , 27.vii. 1985,
leg. J.T. & DA. Polhemus (jtpc, nhmw); 6c? 18 9 apt., 1 9
macr., Palawan, Balsahan Riv., 20 km SW of Puerto Prince-
sa, CL2014, 27.vii.1985, leg. J.T. & D.A. Polhemus (jtpc);
6c? 10 9 apt., Palawan Pr., Busuanga Is., 13 rd.km WNW
Coron, Balulu Falls, 24.2.1996, leg. H.Zettel (81) (zcwa,
gclb, uplb); 2c? 1 9 macr., Palawan Pr., Busuanga Is., 5 km
NW Coron, Mabintangen Riv., 25.-29.2.1996, leg. H.
Zettel (82) (zcwa).
Redescription
Dimensions. - apterous form, length 2.41 (S),
2.50 (9), width 1.66 (cT), 1.74 (9), width of head
1.06 (S), 1.04(9).
Colour (fig. 1 26) . - Whole body prominently yel-
lowish, with distinct dark marks. Eye dark brown or
blackish. Interoculus pale, with one large more or less
round dark mark, which stretches backwards into a
short stripe at each external corner, in most cases very
difficult to recognize as a clear shape; dark mark along
inner margin of eye reaching posterior margin of ver-
tex. Antennal segments black except basal Va of seg-
ment 1. Pronotum with very broad dark stripes so
usually looking prominently dark, with a yellowish
transverse band along posterior margin and one small
yellowish spot at each latero-posterior angle. Mesono-
tum yellowish with dark lateral stripes. Metanotum
yellowish with a triangular black mark at medio-pos-
terior margin. Mesopleura with a broad brownish
stripe, external angles of metacetabula dark (figs. 71).
Fore leg dark with less than basal half yellowish (fig.
92). Middle and hind legs dark. Tergites 1-3 com-
pletely dark, 4-5 dark, with trace of central light
mark, 6-7 prominently yellowish, 8 prominently
167
Tijdschrift voor Entomologie, volume ui, 199s
Fig. 126.
Ventidius (s. str.) wemeri, a-
pterous male, length 2.40 mm.
dark. Laterotergites 1 -2 dark, 3-7 yellowish. Connex-
iva 1-5 dark, 6-7 yellowish. Venter yellowish.
Pilosity. - As a whole, body with longer, more
prominent pilosity. Trichobothria on vertex conspic-
uous. Inner surface of antennal segment 1 with 4 sub-
basal and 1 subapical long spines, dorsal surface with
1 subbasal spine; through basal half of segment 1 to
apical half of segment 4 with fine hair fringe, espe-
cially segments 2-3 more distinct, segments 2-4 with
scattered dark short setae (fig. 64). Dorsum and pleu-
ra bearing long, dark pubescence, more distinct than
usually. Venter clothed by golden pubescence, espe-
cially on mesosternum denser. Fore trochanter and
femur with dark longer trichobothria setae; Inner sur-
face of fore femur and tibia with dark fine hair fringe.
Longer stiff spines scattered along middle and hind
legs. Basal part of middle and hind femora with a very
long trichobothria-like seta on dorsal surface.
Structural characters. - Interoculus broader than
width of an eye, 0.40 : 0.38 in male, 0.41 : 0.37 in fe-
male. Antennae not modified, segment 2 longer than 3
in males (fig. 64), shorter than 3 in females (measure-
ments see table 1). Thorax bulbous, mesonotum some-
what swollen, its lateral width 0.91 (<?), 0.95 (2). In-
tersegmental suture between meso- and metanotum
weak but visible. Metanotum strongly declivent. Low-
er part of metacetabula obtusely triangular, but bilo-
bate in caudal view, internal lobe smaller than external
lobe and curved towards its body, therefore easily ig-
nored (figs. 83, 84). Metasternal tubercle slightly
prominent in male. Fore femur slender and not modi-
fied, claw rising from Vi of segment 2 of fore tarsus.
Anterior margin of abdominal tergi te 1 distinct; anteri-
or margins of tergi tes 2 and 3 obliterated medially and
protruding forward; abdominal sternite 7 as long as the
preceding abdominal sternites together in both sexes.
Male terminalia. - Parameres (fig. 99) symmetri-
cal, slightly curved, not twisted halfway its length,
very slender in middle of its length, apically distinctly
widened, without or with a very faint indication of a
tubercle, with rounded apex; extending beyond geni-
tal segments. Endosoma (fig. 117): dorsal sclerite
long and recurved proximally, ventral sclerite long,
lateral sclerites straight, hooked at two ends, second
lateral sclerites thin, apical sclerite indistinct.
Female terminalia. - Abdominal sternite 7 large,
posterior margin smooth, concave forwards.
Macropterous form. - As apterous form except the
apex of pronotum pointed (measurements see table 3);
length of fore wings from humeri to apex 2.45. colour
pattern of pronotum (fig. 57) black with two irregular
yellowish blotches. Wings anteriorly black with fine
pubescence and with scattered longer black hairs. Pos-
terior membranous part of wings dark brown.
Distribution (map 2). - The Philippines: Palawan
Prov.: Palawan Island, Busuanga Island.
Comparative notes. - This species is very similar to
V. hungerfordi, V. polhemorum sp. n., and V. pilosus
sp. n. For distinguishing characters see under the
comparative notes of these species.
168
Chen & Zettel: Revision ofVentidius
10. Ventidius (s. str.) pilosus sp. n.
(figs. 65, 72, 85, 86, 93, 94, 100, 101, 1 18, 127, map 2)
Type locality. - Indonesia: Sumba.
Type material. — Holotype, a, apt., Indonesia:
Sumba, Nusa Tenggara Timur Prov., Patawang, 55
km E of Waingapu, 15.ix.1991, CL 2603, leg. D.A.
&J.T. Polhemus (jtpc). - Paratypes, 23 6 20 9, apt.,
Id 2 9 macr., same locality data as holotype (jtpc,
nhmw, ppcc); 50* 3 9 apt., Sumba, Nusa Tenggara
Timur Prov., spring and stream 10 km S of Wainga-
pu, 180 m, 13.ix.1991, CL 2597, leg. D.A. & J.T.
Polhemus (jtpc); 53c? 1019 apt., Sumba, Nusa
Tenggara Timur Prov., Mata River, 49 km E of
Waingapu, 15 m, 15.ix.1991, CL 2604, leg. D.A. &
J.T. Polhemus (jtpc, ppcc, nhmw); 41 cT 439 apt.,
Sumbawa, Nusa Tenggara Barat Prov., Bela River, 28
km SW Bima, 100 m, CL 2172, 19.x. 1985, leg. J.T.
& D.A. Polhemus (jtpc, ppcc, nhmw).
Etymology. - Species name refers to the conspicu-
ous pilosity of its body, especially the longer hair fringe
on the ventral surface of fore femur in both sexes.
Description
Dimensions. - Apterous form, length 2.60 (<5),
3.06 (9), width 1.82 (<?), 2.20 (9), width of head
1.15 («Î), 1.26(9).
Colour (fig. 127). - Whole body prominently yel-
lowish, with distinct dark marks. Eye dark brown or
blackish. Interoculus pale, with three longitudinal
brownish marks, one in middle and two along inner
margin of eyes which stretch backwards to the hind
margin of vertex. Antennal segments black except
basal 2/5 of segment 1. Pronotum with very broad
dark stripes so usually appearing prominently dark,
with a yellowish triangular mark along posterior mar-
gin and one small yellowish spot at each latero-poste-
rior angle. Mesonotum yellowish with dark lateral
stripes. Metanotum yellowish with a small triangular
black mark at medio-posterior margin and two large
square dark marks laterally. Mesopleura totally yel-
lowish (fig. 72). Metacetabula with posterior external
angles dark and brownish in anterior external part
(figs. 85, 86). Fore leg dark with basal half of fore fe-
mur yellowish (fig. 93). Middle and hind legs dark.
Tergite 1 completely dark or with two vague yellow-
ish marks laterally, 2-5 dark with triangular central
yellowish marks, 6-7 yellowish, 8 dark. Laterotergites
in some specimens totally yellowish, in some speci-
mens tergites 1-3 dark, 4 prominently dark with cen-
tral yellowish mark, 5-7 yellowish. Connexiva 1-5
dark, 6-7 yellowish. Venter yellowish.
Pilosity. — As a whole, the body with more promi-
nently pilosity. Trichobothria on vertex conspicuous.
Inner surface of antennal segment 1 with 6 subbasal,
Fig. 1 27. Ventidius (s. str.) pilosus, holotype, apterous male,
length 2.60 mm.
and 1 subapical long spines; through basal half of seg-
ment 1 to apical half of segment 4 with dark fine hair
fringe, and with scattered dark short setae (fig. 65).
Dorsum and pleura bearing dark pubescence, more
conspicuous than usually in the genus. Venter
clothed by golden pubescence. Ventral face of fore
trochanter and femur with dark long trichobothrial
setae; in both sexes ventral surface of fore femur with
dark long hair fringe. Longer stiff spines scattered
along middle and hind legs.
Structural characters. - Interoculus broader than
width of an eye, 0.49 : 0.40 in both sexes. Antennal
segment 1 stout but not as broad as in V. hungerfordi,
segment 2 longer than segment 3 in both sexes (fig.
169
Tijdschrift voor Entomologie, volume ui, 1998
65), segment 3 shorter than 4 in male (measurements
see table 1). Mesonotum somewhat swollen, its later-
al width 1.11 (â), 1.27 (?). Intersegmental suture
between meso- and metanotum weak but visible.
Metanotum declivent. Posterior part of metacetabula
obtusely triangular, (figs. 85, 86). Fore femur slender
and not modified, claw rising from Vò of segment 2 of
fore tarsus (figs. 93, 94). Anterior margin of abdomi-
nal tergite 1 distinct; anterior margins of tergites 2
and 3 obliterated medially and protruding forward;
abdominal sternite 7 as long as the preceding abdom-
inal sternites together in both sexes.
Male terminalia. - Parameres (fig. 100, 101) sym-
metrical, slightly curved, distinctly twisted halfway its
length, with slender middle part, apical part distinct-
ly widened, with a faint indication of a tubercle on
ventral external face, apex rounded, extending be-
yond genital segments. Endosoma (fig. 118): dorsal
sclerite long and recurved proximally, ventral sclerite
long, lateral sclerite straight, hooked at two ends, sec-
ond lateral sclerites thin, with third lateral sclerites,
apical sclerite indistinct.
Female terminalia. - Abdominal sternite 7 large,
posterior margin smooth, slightly concave anteriorly.
Macropterous form. - As apterous form except the
apex of pronotum pointed (measurements see table
3); colour pattern of pronotum: black with yellowish
median line and two irregular yellowish blotches.
Wings anteriorly black with fine pubescence. Posteri-
or membranous part of wings dark brown, length of
fore wings from humeri to apex 2.45 mm.
Distribution (map 2). - Indonesia: Nusa Tenggara
Timur Prov. (Sumba Isl.), Nusa Tenggara Barat Prov.
(Sumbawa Isl.).
Comparative notes. - This species is very similar to
V. hungerfordi, V. polhemorum sp. n., and V. werneri,
but V. hungerfordi and V. polhemorum sp. n. are easy to
distinguish by the stouter antennal segment 1 of males
and by the shapes of parameres. The most similar
species is V. werneri from Philippines (Palawan), which
has a similar slender antennal segment 1 and slender
parameres, but the paramere of V. pilosus sp. n. is more
twisted, whereas that of V. werneri is nearly straight;
further V. pilosus sp. n. is larger in body size and length
of middle and hind femora (comp, table 2), has a
denser pilosity of fore femora in both sexes and a longer
antennal segment 2 (compared with segment 3, see
Tab. 1), and differs in colouration of the abdomen.
1 1 . Ventidius (s. str.) modulatus Lundblad
(figs. 59, 66, 73, 74, 87, 95, 102-1 13, 1 19-122,
128, map 3)
Ventidius modulatus Lundblad, 1933: 339-401 (descr., illus-
tr.); Hungerford & Matsuda 1960: 329 (descr., illustr.,
key); Cheng 1965: 162-163 (descr., illustr., key); Kovac
& Yang 1989: 285 (ree); Yang & Kovac 1995: 293 (ree);
Zettel & Chen 1996: 152, 180 (list, ree); Hanboonsong
et al. 1996: 24 (ree).
Ventidius chinai Hungerford & Matsuda, 1960: 331-332,
336 (descr., illustr., key) Syn. n.
Ventidius pubescens Cheng, 1965: 160-161, 163 (descr., il-
lustr., key) Syn. n.
Type locality. - Indonesia: Western Java.
Type material examined. - Ventidius modulatus. Lecto-
type (present designation) c? apt., Indonesia: 'Westjava,
Stausee Tjigombong, South of Buitenzorg, 500 m above see
level, 17.ix.1928' (smnh); paralectotypes: 2 c? apt., from the
same locality (smnh).
Ventidius chinai: Holorype c? apt., Malaysia: 'Malay
Peninsula, Selangore, F.M.S. Kajang Sungei Lang, Feb. 12,
1927', 'C. Dover Coll., F.M.S. Museum' (antennal seg-
ments 3-4 and parts of legs missing) (bmnh); allotype 9
apt., and paratypes 1 <? 1 9 apt from the same locality (gen-
italia of male missing) (bmnh).
Ventidius pubescens. Holotype c? apt., and allotype 9
apt., Malaysia: Johor, Sungai Muar at Rompin, I4.iii.1963,
leg. L. Cheng (bmnh); paratypes: 8(5 39 apt., same locality
(bmnh, zrcs).
Other material examined. - sri lanka: 1 ? apt., 1 c?
macr., Northern Prov., 28 mi. Pullian- kulam Nedunkerni
Rd., 1.-5.V.1966, slow stream, shady corners, leg. P.B.
Karunaratne (jtpc); 26" 2 9 apt., Northern Prov., 127 mi.
Madu Rd., I4.iv.1968, leg. P.B. Karunaratne (jtpc,
nhmw); 2c? 19 macr., North Western Prov., Giriulla
13.X.1957, leg. C.H. Fernando, (semc); 2c? 2 9 Central
Prov., Aruppola, Kandy, 25.vi.1966, edge of river Mahuveli,
leg. P.B. Karunaratne (amnh, jtpc, ppcc); 4c? 59 apt.,
Central Prov., Katagastota, Kalugala, Pinga , Oya,
30.vii.1966, leg. P.B. Karunaratne (jtpc, nhmw); 19 apt.,
2 9 macr., Uva Prov., Helagama, 500 ft., 7--8.xii.1967,
Okkampitiya, from Kumbukkan Oya, leg. P.B.
Karunaratne (jtpc); 3 c? 39 apt., Uva Prov., Aluthnuwara,
29.viii.1967, leg. P.B. Karunaratne (jtpc, nhmw); 2c? 2 9
apt., Uva Prov., Helagama, 500 ft., 7--8.xii.1967,
Meeminna, Heia Pool, at foot of hill, leg. P.B. Karunaratne
(jtpc, ppcc); Thailand: 9 cT 15 9 apt., Chiang Mai Prov.,
Doi Inthanon N.P., Mae Klang Falls, 4.XÌ.1995, leg. H.
Zettel (6) (nhmw, ppcc, nctn); 19c? 249 apt., Chiang Mai
Prov., Huai Hong Khrai, 30 km N. Chiang Mai, stream,
500 m, 31.xi.1994, leg. P.P. Chen & S. Wongsiri (ppcc,
nctn, nhmw); 22c? 149 apt-, Chiang Mai Prov., Fang
Horticultural Station, ponds, 15.xi.1985, CL2201, leg. J.T.
& D.A. Polhemus (jtpc, ppcc, nhmw); 2c? 1 9 macr., Chi-
ang Mai Prov., 7 km NW of Fang, Horticultural Experi-
mental Station, 30.x.-2.xi.l979, Zool. Mus. Copenhagen
Exped. (zmuc); lie? 10 9 apt., 4c? 3 9 macr., Chiang Mai
Prov., stream 10 km NW of Mae Rim, 19.xi.1985, CL
2204, leg. J.T. & D.A. Polhemus (jtpc, nhmw); lc? 5 9
apt., Lampang Prov., Nam Mae Tha, 300 m, loc. 67, shal-
low river, 4.iii.l989, leg. M. Andersen & H. Read (zmuc);
1 9 apt., Lampang, 7 km SW Sop Drop, Mae Nam Wang,
23.iii.1994, A-1041, leg. W.D. Shepard (nhmw); lc? 4 9
macr., Tak Prov., Sam Ngao, at. Bhumiphol Dam, 6.-
8.xi.l979, Zool. Mus. Copenhagen Exped. (zmuc); 1 c? 2 9
Phitsanulok Prov., Subdiv. of Tung Saleangluang N.P., wa-
terfall of Kaek riv., 17.xi.1994, leg. Chen & Piyapichart
(ppcc); 1 c? apt., Phitsanulok Prov., Boi Waterfall, nr. Mae
Nam, Khek River, 300 m, loc. 60, 2.UÌ.1989, leg. Mich. An-
dersen & H. Read (zmuc); 3<? 39 apt. Ubon Ratchan-
170
Chen & Zettel: Revision ofVentidius
Map 3. Ventidius (s. str.) modulatus-
group, partim.
• modulatus
"A- distanti (type locality)
H husbmae (type locality)
ZTTCSp^
thani, Ban Tasala, Maekong River, 17. Nov. 1995, leg. Yupa
Hanboonsong (nhmw); 1 <? apt., Ubon Ratchanthani, Sir-
inton, 16.iii.1996, leg. Yupa Hanboonsong (13) (nhmw);
2 c? 2 9 apt., Ubon Ratchanthani, Tasala, 16.iii.1996, leg.
Yupa Hanboonsong (13) (kkua, nhmw); 3d 4$ apt., same
locality data, 7.VÜ.1996 (18) (kkua, nhmw); 1 9 apt., Ubon
Ratchanthani, Muang, Han Phudua, I6.iii.1996, leg. Yupa
Hanboonsong (14) (kkua); 3c? 3$, all apt., Khon Kaen
Prov., Phu Phan Kham N.P., Ban Noon Huan Chang, Huai
sam Caen, (upper stream of Nam Phong River), N9521,
N9523, 21.xi.1995, leg N. Nieser, (nctn, ppcc); 15c? 8?
apt., same locality, leg. H. Zettel (20a) (nhmw, ppcc); 5 c?
39 apt., same locality, leg. H. Zettel (20b) (nhmw); 3 c?
1 9, all apt., Kanchanaburi, River Kwai, 20.xii.1981, leg. N.
M. Andersen (zmuc); 1 S 5 9 apt., Chantaburi Prov., Hor-
ticulture Research Centre, Dept. of Agriculture, 15 km E of
Chantaburi city, stream, 29.Ì.1995, leg. P.P. Chen (ppcc,
nhmw); 1 c? 19 apt., Surin Prov., Surin, 150 m, 5.-
10.12.1995, leg. P. Schwendinger (nhmw); 9c?, 29 apt.,
Songkhla Prov., Hat Yai, SW of Songkhla, 21.xi.1979, leg.
P. Nielsen (zmuc, nhmw); viet nam: : 4c? 39 QuangTri,
1 mi. N of Quang Tri, 9.v., 15.v., and 9.vi.l970, leg. A.R.
Gillogly (jtpc, nhmw); 5 c? 79 apt., Gia Lai-Kontum, 20
km N of Pleiku, 650 m, 9.V.1960, leg. S. Quate (bimc, jtpc,
nhmw); 2c? 5 9 apt., Gia Lai-Kontum, Pleiku, 700 m, 8.-
14.V.1960, leg. S. Quate (bimc); 1 9 apt, Gia Lai-Kontum,
25 km SW of Pleiku, 400 m, 12.V.1960, leg. S. Quate,
(bimc); Lam Dong, 45 km E of Da Lat, 850 m, 5.V.1960,
leg. S. Quate, 2 c? (a) (bimc); 5 c? 8 9 apt., 2c? 49 macr.,
Song Be: Nam Cat Tien NP, 1.-15.5.1994, leg. Pacholatko
&C Dembicky (nhmw, ppcc); Malaysia: 12c? 13 9 apt., Pa-
hang, pond 54 km SW of Kuantan, CL 2085, 22.viii.1985,
leg. J.T. & DA. Polhemus (jtpc, nhmw); 32c? 50 9 apt.,
Pahang, river 65 km NE of Segamet, CL 2086, 22.viii.1985,
leg. J.T. & DA. Polhemus (jtpc, nhmw, ppcc); 2 9 apt.,
Pahang, King Geo. V. Nat. Park, Kuala Tahan, 12.-
15.xii.1958, leg. T.C. Maa (bimc); 19 apt., Pahang, S.
Kinchin, 13.vi.1989, leg. CM. Yang, zrcs2617 (zrcs); lc?
1 9 apt., Johor, swamp forest stream, 15 km W of Sedili Be-
sar, 20 m, 16.X.1986, CL 2218, leg. J.T. & DA. Polhemus
(jtpc); 1 c?, 1 9 apt., 2c? macr., Johor, 27 km S of Mersing,
slow shaded stream, CL 2058, I4.viii.1985, leg. J.T. & D.A.
Polhemus (jtpc); 1 c? 4 9 apt., Johor, blackwater river 2 km
W of Sedili Besar, 0-50 m, 1 6.x. 1986, CL 2219, leg. J.T. &
D.A. Polhemus (jtpc, nhmw); 7c? apt., Johor, Tg. Sedili,
Sg. Selang, muddy water, sandy bottom, up to 1.5 feet wa-
ter, 22.iv.1992, Y792A, leg. K.L. Yeo (zrcs); 4c? 49 apt.,
Johor, S. Bong, 5.iv.l992, Y788L, leg. K.L. Yeo (zrcs); 8c?
59 apt., Johor, S. Endau, nr. base camp (partly night col-
lecting), 3.-5.iv.l992, Y788A, Y788B, and Y788J, leg. K.L.
Yeo (zrcs, nhmw); lc? 49 apt., Johor, Endau, Sungai
Jasin, 3.iv.l992, L185, leg. H.K. Lua (zrcs); lc? 1 9 apt.,
Johor, S. Jenahang, 4. via. 1963 (zrcs); 19 macr., Negeri
Sembilan, 10 km S of Seramban, CL 2059, l4.viii.1985,
leg. J.T. & D.A. Polhemus (jtpc); 2<? 3 9 apt., Negeri Sem-
bilan, 9 km S Simpang Pertang, Serting River, 10. iv. 1994,
A-1062, leg. W.D. Shepard (ess, nhmw, ppcc); 28c? 269
apt., Negeri Sembilan, 14 km NW Kota, unnamed stream,
9.iv.l994, A-1056, leg. W.D. Shepard (ess, nhmw, ppcc);
1 9 apt., Negeri Sembilan, Ulu Bahau, Rubber Estate,
17.iii.1964, zrcs6.6481 (zrcs); lc? apt., Negeri Sembilan,
S. Jelai, l.xi.1963 (zrcs); 2c? apt., Selangor, Dusun Jua, 17
miles from Diteh, 26.viii.1926, coll. Dover (bmnh); 4c? 29
apt., Selangor, North Selangor Peat swamp-forest stream at
43 km mark on road to Sg. Besar, 19.vi.1991, leg. H.K. Lua
& Mala, LHK159 (zrcs, nhmw); 15 9 apt., Selangor, Am-
pang, nr. Sg. Kongsi Laban, 500, small pond, 13.iii.1964,
leg. L. Cheng, zrcs6.6456-6470 (zrcs); 2 9 apt., Upper
Perak, Longgong, surface of irrigation channel, fast water,
22.X.-5.XÌ.1926, coll. Dover (bmnh); 4c? 49 apt., Kelantan,
171
Tijdschrift voor Entomologie, volume 141, 1998
Sg. Sat, ll.xii.1964, leg. A. Jothy (jtpc, nhmw); 1 â apt.,
Kelantan, Sungai Hulu Besut, 6 km (W?) after Kampang
Keruah, on road to hydroelectric station, 20.iii.l992, LI 84,
leg. H.K. Lua (zrcs); 4S 49 apt., P. Tioman, tributary of
Sg. Paya, brackish water, 25.vi.1996, leg. CM. Yang, ' Yii7
(zrcs, nhmw); 4â 4? apt., P. Tioman, Sg. Kililing,
27.vi.1996, leg. H.K. Lua, Y136 (zrcs); IcT 19 apt.,
Terengganu, Sg. Terengganu tributary, Sekayu, 16.v. 1995,
leg. Bruce Tan &C Sumita, TG6 (zrcs); 4 c? apt. Prov.?, Sun-
gai Kingsi, Laban, leg. L. Cheng (jtpc); 30* 39 apt., Prov.?,
Sg. Belat, 26 km from Kuan tan, 15. v. 1995, leg. Bruce Tan
& Sumita, TG3 (zrcs, nhmw); Singapore: 2$ apt., 16.5
mis. from Mersing, Mersing-Kluang Rd., 4.viii.l963, leg. L.
Cheng (bmnh); 6â 5? apt., pond nr. MacRitchie Reser-
voir, 11.5.1993, NS25 (zrcs, nhmw); 69 apt, Botanical
Gardens pond, 6.iv.l990, leg. CM. Yang & H.K. Lua
(zrcs); 46 apt., Bukit Batok Nature Park, 10.7.1995, leg.
H.K. Lua, LHK282 (zrcs); 43 3 9 , stream at Lower Pierce,
21. vii. 1990, Y21B (zrcs); 29 apt., Stream H, Sime Road,
5.vi.l995, leg. H.K. Lua et al. NS181 (zrcs); Indonesia:
27 S 189 apt., \6 19 macr., Sumatra, Bengkulu Prov.,
Bantiring River, 7 km E of Bengkulu, 30 m, 6.ix.l991, CL
2580, leg. D.A. & J.T. Polhemus (jtpc, nhmw, ppcc); lo*
apt., (Borneo), Kalimantan Timur Prov., waterfall 4 km S of
Kota Bangun, 29.viii.1985, CL 2095, leg. J.T. & D.A. Pol-
hemus (jtpc); 1 S apt., 1 9 macr., Kalimantan Timur Prov.,
Borneo, waterfall and stream, 1 1 km NE of Samarinda, CL
2091, 27.viii.1985, leg. J.T. & D.A. Polhemus (jtpc); 5 c?
79 apt., 19 macr., 8 larvae, Kalimantan Timur Prov.,
Kedang R., 12 km above Maura Kedang, 0.6°S, 116.2°E,
leg. M.S. Christensen (jTpc, ppcc, nhmw). 10 males macr.
Myanmar (Burma): Bago Division, 33 km W of Oktwin,
Bago Yoma, Sein Yai Forest Camp, Selnyay River, 170 m,
29.x. 1998, leg. H. Schillhammer (nmw, ppcc).
Redescription
Dimensions. - Apterous form. Length 2.54 (<S),
3.10 (5), width 1.90 (<?), 2.05 (9), width of head
1.10 (â), 1.10(C).
Colour (fig. 128). - Whole body prominently yel-
lowish, with distinct dark marks. Eye blackish. Inte-
roculus pale, with one large dark mark, dark mark
along inner margin of eye reaching posterior margin of
vertex, but in the case of paralectotypes indistinct. An-
tennal segments dark, except basal Vò-Va of segment 1.
Pronotum usually yellowish with two lateral dark
marks. Mesonotum yellowish with dark lateral stripes.
Metanotum with square lateral dark mark. Mesopleu-
ra yellowish, its anterior half with a dark stripe; exter-
nal angle of metacetabula dark (figs. 73, 74). Fore leg
dark with basal V3 of fore femur yellowish (fig. 95).
Middle and hind legs dark. Tergite 1 completely dark,
2-5 dark, with central light mark, 6-8 yellowish. Lat-
erotergites 1-2 dark along the edge, 5-7 yellowish,
Connexiva 1-5 dark, 6-7 yellowish. Venter yellowish.
Pilosity. - Inner surface of antennal segment 1 with
3 subbasal, and 1 subapical long spines; through basal
half of segment 1 to apical half of segment 4 with sil-
very fine hair fringe; segments 2-4 with scattered dark
short setae (fig. 66). Dorsum and pleura bearing dark
pubescence. Venter clothed by golden pubescence,
especially on genital segments, the pilosity very long
and dense. Fore femur and tibia with silvery fine hair
fringe. Longer stiff spines scattered along middle and
hind legs. Basal part of middle and hind femur with a
very long trichobothria-like seta on dorsal surface.
Structural characters. - Interoculus broader than
width of an eye, 0.42 : 0.31 in male. Antennae not
modified, segment 2 longer than 3 in males (fig. 66),
roughly of same length in females (measurements see
table 1). Thorax not bulbous, mesonotum somewhat
swollen, its lateral width 1.02. Intersegmental suture
between meso- and metanotum distinct. Metanotum
somewhat declivent. Lower part of metacetabula trun-
cate (fig. 87). Metasternal tubercle slightly prominent
in male. Fore femur slender and not modified, claw
rising from Vi of segment 2 of fore tarsus. Anterior
margin of abdominal tergite 1 distinct; anterior mar-
gins of tergites 2 and 3 obliterated medially and pro-
truding forward; abdominal sternite 7 as long as the
preceding abdominal sternites together in both sexes.
Male terminalia. - Parameres (figs. 102-113) sym-
metrical, twisted in middle part, apically strongly
widened, with distinct tubercle, with rounded apex,
extending beyond genital segments. Endosoma (fig.
1 19-122): dorsal sclerite long and recurved proximal-
ly, ventral sclerites long, lateral sclerites straight, prox-
imal parts broadened and hooked, second lateral scle-
rites thin, apical sclerite indistinct.
Female terminalia. - Abdominal sternite 7 large,
posterior margin smooth, slightly concave anteriorly
(fig. 59).
Macropterous form. - (After specimens from Sri
Lanka). Colouration: pronotum black with a large
round yellowish central blotch. Wings anteriorly black
with fine pubescence and with scattered longer black
hairs. Posterior membranous part of wings dark
brown. The other morphological characters as in
apterous form except the apex of pronotum pointed
(measurements see table 3); length of fore wings from
humeri to apex 2.02.
Variability. - In contrast to most other species of
Ventidius, V. modulatus shows a surprising variability
in dorsal colour pattern and size, but specimens from
the same sample are usually similar in these charac-
ters. Bright yellowish specimens are mainly found in
Malaysia, where dark-coloured populations are also
found. In a large sample from Negeri Sembilan (A-
1056, leg. Shepard) all specimens have a completely
yellowish thorax. Specimens from Sri Lanka always
have dark lateral stripes on mesonotum, specimens
from Thailand mostly so; there are only a few samples
with differently coloured specimens, e.g. that from
Udon Thani Prov., Thailand. Pubescence of thorax is
also similar between specimens of the same sample.
We include within V. modulatus also a dark-
coloured population from Viet Nam (Gia Lai-Kon-
172
Chen & Zettel: Revision ofVentidius
Fig. 128. Ventidius (s. str.) modulatus, lectotype, apterous
male, length 2.54 mm.
turn), which shows a slightly thickened antennal seg-
ment 1 in males (length : width = 12.5-13.5), denser
and longer pilosity and a more flattened body than in
other populations within the large area of V. modula-
tus. These specimens resemble somewhat V. hunger-
fordi, but, beside genitalia structures, the antennal
segment 1 of males is distinctly less stout (9.5-1 1.5 in
V. hungerfordi) and the pleural colour pattern is dif-
ferent. We have also seen intermediate V. modulatus
specimens from Northeast Thailand, so that we are
sure that the Viet Nam population is no more than a
geographic, infrasubspecific form of V. modulatus.
Distribution (map 3). - Sri Lanka: Northern Prov.,
North Western Prov., Central Prov., Uva Prov.; India:
West Bengal State; Thailand: Chiang Mai, Lampang,
Tak, Phitsanulok, Khon Kaen, Ubon Ratchanthani,
Kanchanaburi, Chantaburi, Surin, Songkhla; Viet
Nam: Quang Tri, Già Lai-Kontum, Lam Dong, Song
Be; Malaysia: Selangor, Johor, Perak, Pahang, Negeri
Sembilan; Singapore; Indonesia: Sumatra, Java, Kali-
mantan.
Comparative notes. - This species is similar to the
four preceding species (V. hungerfordi, V. polhemo-
rum sp. n., V. werneri, and V. pilosus sp. n.), and dif-
fers mainly in the slender antennal segment 1 of
males (much thicker in V. hungerfordi and V. polhe-
morum, slightly thicker in V. werneri and V. pilosus),
and a shorter pubescence on metapleura. The para-
mere is especially similar to that of V. hungerfordi,
but differs from that of the other species by having a
distinct tubercle.
Remarks. — As the synonymies of the taxa V. mod-
ulatus, V. chinai, and V. pubescens are problematical,
we feel obliged to give a detailed analysis of the type
specimens:
We have studied three of the four male syn types of
V. modulatus Lundblad 1933, from West Java, which
are deposited in the smnh. One of them bears the la-
bel 'Typus' and is presently designated and labelled as
lectotype, the other two with the label 'Paratypus' as
paralectotypes.
Ventidius chinai was based on two males and two
females from Selangor in Malaya. In the comparative
notes by Hungerford & Matsuda (1960) it is com-
pared with V. aquarius, but this species is not closely
related. In the key by Hungerford & Matsuda (1960)
V. chinai is compared with V. werneri and V. henryi by
colouration; due to a mistake (paragraph 6'. is miss-
ing) the comparison with V. modulatus is not clear,
but mainly based on the shape of parameres and more
(in V. modulatus) or less (in V. chinai) developed su-
tures between the meso- and metanotum and the
metanotum and tergite 1. The description of the para-
mere of V. chinai is based on the incomplete paramere
of the holotype (apical part lacking) and therefore use-
less for a comparison; strangely enough we have seen
two males from Selangor (also from the Dover collec-
tion) with similarly broken parameres. We could not
find any proper differences in the development of tho-
racic sutures between V. chinai and V. modulatus. Dif-
ferences mentioned by Hungerford & Matsuda
(1960) may be due to the figure of Lundblad (1933:
pi. xii) which shows very pronounced sutures; in fact
the sutures are faint but partly dark-coloured in one
type specimen investigated. The dark colour pattern is
extremely reduced in the types of V. chinai. The tho-
racic hairs are partly rubbed off, so that an investiga-
tion is difficult. In the centre of the mesopleura the
hairs are short as in the types of V. modulatus.
Ventidius pubescens was based on numerous speci-
mens from Johor in Malaya (Cheng 1965). We have
studied the holotype and several paratypes, which
have a uniformly reduced dark colour pattern. Cheng
(1965) re-described V. modulatus after 'a mating pair'
from Pahang, Malaysia, but compared V. pubescens
with V. chinai on the assumption that '[t]he colour
pattern is also quite a constant character' (Cheng
1965: 153). The main difference between V. pubescens
and V. chinai is, according to Cheng (1965), the dif-
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Tijdschrift voor Entomologie, volume i4i, 1998
ferent shape of parameres. As this is based on the bro-
ken paramere of V. chinai, this character is useless.
Furthermore, Cheng (1965) stated: ' V. pubesce ns also
differs from the latter species [= V. chinai] in the pos-
session of the mesosternal groove, densely haired body
and differently proportioned antennal and leg seg-
ments.' The 'mesosternal groove' mainly consists of an
internal ridge medianly on the mesosternum and can
be more or less seen through the cuticula; it is more
easily visible in specimens preserved in alcohol ( V. pu-
bescens paratypes) than in dried specimens (as type
specimens of V. chinai). We could not find any im-
portant differences in ratios of the antennal and leg
segment of the type specimens of V. chinai and V. pu-
bescens for these parts which are still present in the
types of V. chinai. Indeed there is a difference in the
pubescence of V. pubescens and V. chinai (and V. mod-
ulatili), not so much in the general dorsal appearance
(which we think is due to the bad condition of the
type specimens of V. chinai), but more in a longer pu-
bescence in the middle part of mesopleura of V. pubes-
cens. This character can be seen only under high mag-
nifications and is the only character we found to
distinguish V. pubescens from V. modulatus and V. chi-
nai. Body pubescence has been shown to be important
for protection against UV-radiation in different gen-
era of Gerridae (Cheng et al. 1978, Andersen 1990);
therefore it could be variable under different micro-
habitat conditions. As the types of V. pubescens corre-
spond in all other characters either with V. modulatus
and/or with V. chinai, we regard it as synonymous
with V. modulatus. Although there may be some
doubts about the synonymy of V. pubescens and V.
modulatus, we regard it presently more practical to
treat them as one species than to separate them only by
the difference in pubescence which is very difficult to
observe. Recently, a sample including only macropter-
ous specimens was collected at Bago Yoma, Burma (c.
18°47'N, 96°21'E), whcih is close to the type locality
of V. distanti (c. 20°35'N, 96°58'E). The paramere
shows the typical shape of the V. modulatus-group.
We suspect that these macropterous specimens from
Bago Yoma are V. distanti. In that case, further study
of V. distanti and V. modulatus will be needed (see also
under V. distantì).
12. Ventidius (s. str.) distanti Paiva
(fig. 129, map 3)
Ventidius distanti Paiva, 1918: 25, pi. VIII, fig. 4 (descr., il-
lustr.); Esaki 1928: 511 (comparative notes); Dover
1929: 69 (misidentif.); Esaki 1930: 18 (considers as valid
species, misidentif.); Lundblad 1933: 372 (list, comp,
with modulatus); Hungerford & Matsuda 1960: 332 (dis-
cussion, key).
Type locality. - Myanmar (burma): Yawnghwe State.
Types [not examined]. — 'Described from several speci-
mens in alcohol, from the top of gorge of the He-Ho River,
Yawnghwe State, ca. 3,500 ft., 7-iii-1917' (Paiva 1918).
Description
Since we have no material either from the type se-
ries or other specimens from the type locality, we
adopt the main description points given by Paiva
(1918) which are based mainly on colour characters.
Dimensions. — Apterous form. Length 3.00.
Colour and structural characters. - 'Head black
with a large patch at base, and a transverse fascia at
apex of face yellowish ochraceous; eyes silvery grey,
with a black patch on the disk; antennae black, basal
half of first joint yellowish. Pronotum very short,
black, a narrow ochraceous weaved fascia at basal
margin, anterior margin slightly concave, posterior
margin almost straight. Mesonotum large, about as
long as its greatest breadth (Esaki (1930) pointed out
that '[h]is "mesonotum' evidently includes a part of
the metanotum'), covered with decumbent hairs, disk
obliquely striate on anterior area, ochraceous, with
two broad lateral black fasciae curved inwards anteri-
orly and meeting narrowly on anterior margin, each
extended posteriorly to meet a curved fascia on the in-
termediate acetabula; a large subtriangular patch at
centre of posterior margin; the posterior lateral angles
narrowly dull black. Metanotum dull black with a
small ochraceous spot near each basal angle. Ab-
domen above dull white, the basal segment, a spot at
lateral margin of each segment and the apical segment
black. Underside pale ochraceous; legs black, base of
anterior femora ochraceous.'
Distribution (map 3). - Myanmar: Yawnghwe.
Remarks. - The type material of this species was
deposited in the collection of the Zoological Survey
of India (type No. 7152/H.i.), but is so far not avail-
able for study. We quote here the notes from the revi-
sionary paper of Hungerford & Matsuda (1960): 'Dr.
P. Kapur of the Calcutta Museum is now unable to
find this type in their museum and it may have been
lost by high water in the temporary quarters in 1943.
This is most unfortunate since only a re-examination
or the type of a study of a series of specimens from the
type locality the identity of Ventidius distanti can be
established.'
Judging from the drawing (fig. 129) and the de-
scription given by Paiva ( 1 9 1 8) we consider V. distan-
ti as belonging to the V. modulatus-group; the speci-
men figured is probably a female. Species of the
subgenus Ventidioides usually have the pronotum
completely black (never with a yellow mark medially
on hind margin as figured by Paiva). Species of the V.
aquarius-gwup are larger in size (Paiva states 3 mm for
V. distanti) and have longer antennae and stouter mid-
dle and hind femora than figured by Paiva (1918). Re-
174
Chen & Zettel: Revision ofVentidius
Fig. 129. Ventidius (s. str.) distanti, apterous male, repro-
duced from Paiva's drawing (1918).
cently we have obtained records of V. modulatus from
India and Sri Lanka, which shows that this species is
widely distributed and probably also occurs in Myan-
mar (although we have no records from this country).
As the description given by Paiva (1918) is not suf-
ficient, there was a confusion about the species named
V. distanti during the following decades. Dover
( 1 929) recorded it from Malaya (Selangor) and added
to the confusion; his material was considered as be-
longing to two different species (see Hungerford &
Matsuda 1960). One of them was established as a new
species, V. malayensis, by Hungerford and Matsuda
(I960), while the other smaller specimen-which was
identified as V. distanti also by Esaki and China-is a
Ventidioides sp. We agree with Hungerford & Matsu-
da ( 1 960) that the morphological characteristics show
clearly that this specimen belongs to the subgenus
Ventidioides, and reject the conspecific status of this
specimen with V. distanti. The only Ventidioides
species presently known from Malaya (and from Se-
langor) is V. pulai.
Lundblad (1933) stated that his new species, V.
modulatus, was similar to V. distanti, but differed in
the length of the antennal segments: segment 3 was
longer than segment 2 in V. distanti, but shorter than
segment 2 in K modulatus. In this case-because he
had only four males at hand-Lundblad (1933) walked
right into a deep trap: he could not know that in this
species complex ( V. modulatus-gïoup) usually the ra-
tio between lengths of antennal segment 2 and 3 is
completely different in males and females. If we are
right that Paiva (1918) figured a female, indeed there
is no obvious difference between V. distanti and V.
modulatus. Furthermore, Lundblad (1933) trusted
the paper of Esaki (1930): Esaki had a male at hand of
which the antennal segment 2 is also shorter than seg-
ment 3; but this specimen even belongs to another
subgenus (see above).
A recent sample from Burma (see also under V.
modulatus) including only macropterous specimens,
was collected at Bago Yoma (c. 18°47'N, 96°21'E),
which is close to the type locality of V. distanti (c.
20°35'N, 96°58'E). It is quite possible that these
macropterous specimens from Bago Yoma areK dis-
tanti. A definitive conclusion is difficult to draw with-
out checking the type material.
Finally, we are of the opinion that V. distanti should
be a valid species of the genus Ventidius within the V.
modulatus-group. It cannot be V. henry i because of the
colour pattern figured by Paiva (1918) (see fig. 129),
and probably not V. hungerfordi which has a thick an-
tennal segment 1 and is so far only known from
Southeast Asia. But until material from the type local-
ity is available, we consider it premature to make V.
distanti a senior synonym of one of the known sibling
species in this group.
13. Ventidius (s. str.) sushmae Gupta nomen
inquirendum
Ventidius sushmae Gupta, 1981: 99-102, figs. 1-12 (descr., il-
lustr.).
Type locality. - india: West Bengal, Darjeeling, Sukna.
Types [not available for examination] . - 'The type speci-
mens are for the time being retained in the department of
Zoology, B.S.A. College, Mathura (U.P.) but subsequently
would be deposited in the National Collection of Zoological
Survey of India, Calcutta' (Gupta, 1981). The type serie in-
cludes 10 apterous males and 8 females, and 6 macropterous
males and 4 females.
Description
We were not able to study type material of V. sush-
mae. We adopt the description given by Gupta (1981)
as below:
Dimension. - Apterous form, length 2.40 (a),
2.60 (9), width 1.85 (<?), 2.10 (9), width of head
1.05 (<?), 0.95 (9).
Colour. - Dorsal surface yellowish-brown in
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Tijdschrift voor Entomologie, volume i4i, 1998
ground colour with black markings. Head yellowish-
brown, a median somewhat triangular and a pair of
lateral black stripes, confluent anteriorly. Pronotum
yellowish-brown with lateral longitudinal black
stripes. Mesonotum yellowish-brown with a pair of
lateral longitudinal black stripes. Metanotum yellow-
ish-brown with black stripes on fifth, sixth and sev-
enth tergites. Venter light brown. Connexivum yel-
lowish-brown laterally. Antennae and legs brown.
Pilosity. - Fore femur of male not clothed with
long hairs on inner margin.
Structural characters. - (The measurement of an-
tennnae and legs were given by Gupta for both apter-
ous and macropterous form. As the scale was not in
millimetres, his data are excluded from the tables in
this paper). Head including eyes much wider than
long. Eyes exserted, rounded on outer margin,
strongly emarginate on inner half margin, covering
entire lateral margin of pronotum and anterior angle
of mesonotum. Antenniferous tubercles not devel-
oped. Proportional length of antennal segments see
table 1. Clypeus with balsa margin lost, mandibular
and maxillary plates well demarcated from each oth-
er. Rostrum short, clearly surpassing hind margin of
prosternum, third segment about three times longer
than last segment [7.2: 2.4 (cT), and 7.5: 2.5 ($)].
Pronotum short, transverse, lateral margin rounded,
both anterior and posterior margins concave. Inter-
segmental suture between mesonotum and metan-
otum poorly defined dorsally, obliterated laterally in
front of metathoracic spiracle, mesosternum without
a small tubercle on median longitudinal axis. Metan-
otum without median longitudinal sulcus, lateral lon-
gitudinal suture not meeting intersegmental suture.
Metacetabular suture clearly reaching anterolateral
angle of first abdominal tergite dorsally. Metacetabu-
lum broad, with posterolateral angle simple.
Metasternum highly reduced, represented by a small
transverse subtriangular plate. Omphalium highly re-
duced, located closer to posterior margin than to an-
terior margin of metasternum. Omphalial groove ab-
sent. Fore leg relatively long, femur slender, simple,
without a tubercle in male; tibia with a conspicuous
narrow apical processus; tarsus with first segment
much shorter than second segment. Claws arising
from about middle of second segment. Abdomen
broad and short. First tergite nearly straight on ante-
rior margin. Anterior margin of second and third ter-
gites produced anteriorly, obliterated medially. Ab-
dominal spiracles hidden beneath metacetabula and
hind legs, situated about in the middle between both
the margins of each segment. Ventral longitudinal su-
ture of connexivum not retained. Median ventral lon-
gitudinal carina absent. Seventh ventrite a little short-
er than all preceding ventrites together, deeply
concave on apical margin.
Male terminalia. - Eighth segment with dorsal pos-
terior margin broadly rounded, ventrally concave on
apical margin. Ninth segment with suranal plate sim-
ple; pygophore with apical margin simply rounded;
parameres well developed, symmetrical. Endosoma
with definitive dorsal plate curved back along apical
margin of endosoma; simple at apex; ventral plate in-
distinguishably fused with dorsal plate, supported by
basal plate; lateral plates simple, elongated, weakly
sclerotized, with small downward projection apically.
A V-shaped sclerite lying over dorsal plate.
Female terminalia. - Seventh segment ventrally
distinctly longer than all preceding segments togeth-
er, nearly straight on hind margin, narrowly rounded
apically; intervalvular membrane with its apical mar-
gin broadly concave, valva large totally membranous.
Macropterous form. - Length 3.58 (S), 3.72 (9),
width 1.90 (cT), 1.92 (5), width of head 1.10 (<J),
1.14(9).
Wing venation: Hemelytra with distinct emboli-
um. R+M+Cu vein divided into two veins, R+M and
Cu at basal third of the wing; two apical cells are
formed, lower cell is relatively longer than vein A
joined at its extremity. Vein also joined with rear
margin of wing at about middle by a short cross vein.
Distribution (map 3). - India: West Bengal.
Remarks. - Gupta (1981) correctly put this species
in subgenus Ventidius (s. str.). He discussed and
keyed the difference of the two subgenera: Ventidius
(s. str.) and Ventidius (Ventidioides) . The comparative
notes of this species with other species of V. modula-
fttf-group were not given. But he mentioned that he
has compared this species with V. aquarius Distant,
although he wrote: 'Unfortunately, the type specimen
(of V. aquarius Distant at Z.S. I., Calcutta) is poorly
mounted on a slide and is damaged.' The illustration
given by him is reasonable and shows great similarity
to V. modulatus Lundblad. The original description
and drawings, especially the lateral view of paramere,
fit very well with V. modulatus. Further, we have stud-
ied specimens of V. modulatus from Sri Lanka. Al-
though we suspect strongly its status as a good
species, until checking the type material we treat here
this species as a valid species in the V. modulatus-
group of the subgenus Ventidius.
Ventidius ( Ventidioides) Hungerford & Matsuda
Ventidius (Ventidioides) Hungerford & Matsuda, 1960:
335-336. Type species by monotypy: Ventidius {Ventid-
ioides) kuiterti Hungerford & Matsuda.
Diagnosis. - Dorsal dark colouration more promi-
nent and not varying as much as in Ventidius s. str.;
antennae stouter than in Ventidius s. str., in males the
length of antennal segment 1 approximately as long
176
Chen & Zettel: Revision ofVentidius
as segments 2-4 together, from basal half of segment
1 to segment 3 or 4 with a fine pubescence of erect
hairs ventrally; posterolateral angle of metacetabula
bilobate; mesosternum of male with a small, in some
species very prominent, tubercle at the end of the me-
dian groove, either with a row of dark setae anteriorly
of the tubercle, or the tubercle more or less covered
with a tiny tuft of setae; surface of male fore femur in
middle of its length with a small tooth, a broad tu-
bercle or simple; parameres asymmetrical, left para-
mere stronger than right one.
Distribution (maps 4, 5). - Widely distributed in
the Oriental region (in the Southeast Asian mainland
from Myanmar to Malaysia, and in Borneo) and
reaching the Wallacea (Sulawesi) . Unknown from the
Indian region.
Identifications. - The following key for apterous
males may be also used for macropterous males in
most characters, but as the thickening of fore femur
and fore tibia is less developed in this morph, this
character can not be used to distinguish V. kurtokala-
mi and V. nieseri sp. n. Females are very difficult to
distinguish in most species, but females of the V.
kuiterti-group differ partly in colour and length of an-
tennal segments, body length and colouration. The
Sulawesian species of the V. xiphibion-gxowp (V.
xipbibion and V. xyele) also show a different colour
pattern in females, which is not very reliable, because
both species show some variations. The same is true
for the Bornean species V. kurtokalami and V. nieseri
sp. n. The female of V. heissi sp. n. is unknown.
Fig. 130. Ventidius (Ventidioides) kuiterti, paratype, apterous
male, length 2.46 mm.
Key to apterous males of the subgenus Ventidioides
1 . Ventral surface of fore femur with a small, sharp
tooth in middle of its length (fig. 148); mesoster-
num without a row of dark setae anterior of tu-
bercle, but the tubercle may be more or less cov-
ered by dark setae (fig. 132); endosoma with two
pairs of lateral sclerites (fig. 4) (V. kuiterti-group;
species from the Southeast Asian mainland) 2
- Ventral surface of fore femur without a sharp
tooth, but sometimes in middle of its length with
a blunt tubercle or thickening (figs. 198-202);
mesosternum with a row of black setae anterior of
tubercle (fig. 215); endosoma with three pairs of
lateral sclerites (fig. 5) (V. xiphibion-gronp; species
from Borneo and Sulawesi) 5
2. Antenna! segment 3 shorter than segment 4, seg-
ment 3 and 4 partly yellowish 3
- Antennal segment 3 subequal to segment 4, seg-
ments 3 and 4 uniformly brown 4
3. Body length less than 2.6 mm; yellowish marks of
meso- and metanotum separated by a black stripe
(fig. 130); antennal segment 4 slightly longer
than segment 3; yellowish mark on antennal seg-
ment 4 less distinct; parameres as in figs. 167,
168 (Myanmar, NE India) V. kuiterti
- Body length more than 2.6 mm; yellowish marks
of meso- and metanotum partly confluent (fig.
183); antennal segment 4 distinctly longer than
segment 3; yellowish mark on antennal segment
4 very distinct; parameres as in figs. 169, 170
(Thailand, Vietnam) V. karen
4. Left paramere in distal part long and slender, not
twisted (fig. 176); right paramere small and
rather slender (fig. 175); segment 8 with densely
developed long dark pilosity (fig. 182) (Thailand)
V. lundbladi
- Left paramere in distal part plate-like, apically
twisted (figs. 172, 174); right paramere slightly
broader (fig. 171, 173); pilosity of segment 8 less
developed (Thailand, Laos, Malaya) V. pulai
5. Fore trochanter without a tuft of black setae; left
and right parameres hooked upward halfway
their length, with slender base (figs. 223, 224,
227, 228) (Bornean species) 6
- Fore trochanter with a distinct tuft of black setae
(fig. 213); inner surface of fore femur and fore
177
Tijdschrift voor Entomologie, volume i4i, 1998
Figs. 131-132.
Ventidius karen. — 131, dorsal
view of macropterous male
(appendages removed, after
Lansbury); 132, ventral view
of apterous male (genital seg-
ments removed, after Lans-
bury).
Fig. 133.
V. pulai, pronotum of macro-
pterous form.
Fig. 134.
V. lundbladi, lateral view of
head, showing the tricho-
bothria of interoculus.
132
tibia without a swelling halfway its length, but
slightly broadened (figs. 198, 199); left: and right
parameres hooked upward close to the strongly
enlarged base, left paramere distinctly broader
than right one (figs. 219-222) (Sulawesian
species) 8
Inner surface of fore femur and fore tibia with a
more or less prominent swelling halfway its
length (figs. 200, 201); row of black setae on
mesosternum long; left and right parameres slen-
der and pointed apically 7
Inner surface of fore femur and fore tibia without
distinct swelling halfway their length (fig. 202);
row of black setae on mesosternum less devel-
oped; both parameres thick, left paramere curved
apically, right paramere blunt (figs. 225, 226) ....
V. heissi
Brighter species, yellowish marks on metanotum
large, metacetabulum with two separated black
marks (figs. 207, 208); thickening of fore tibia
stronger developed; along mesosternal groove with
a row of short dark setae; endosomal apical acces-
sory sclerite indistinct (fig. 232)
V. kurtokalami
Darker species, yellowish marks on metanotum
small (fig. 236), metacetabulum in most speci-
mens with confluent black marks (figs. 209, 210);
thickening of fore tibia mainly by erect pilosity;
along mesosternal groove with a row of long black
setae (fig. 215); endosomal apical accessory scle-
rite distinct (fig. 233) V. nieseri
colouration darker, yellowish marks on metan-
otum usually separated from that of mesonotum;
mesopleura with a dark long stripe running
through its length; left and right parameres with
elongate tips, right paramere not much shorter
than left one (figs. 221, 222); endosomal apical
accessory sclerite indistinct (fig. 231) V. xyele
colouration brighter, yellowish marks on metan-
otum usually not separated from that of mesono-
tum; mesopleura with a small dark mark anteri-
orly, Left paramere with acute, but not elongate
178
Chen & Zettel: Revision ofVentidius
tip (fig. 219), right paramere with less slender tip
and much shorter than left one (fig. 220), endo-
somal apical accessory sclerite distinct (fig. 230) .
V. xiphibion
The Ventidius kuiterti-group
Diagnosis. - Male fore femur in middle of its length
with a small distinct ventral tooth; at the end of medi-
an mesosternal groove with a small prominent tuber-
cle covered with short setae; parameres with blunt
apex (except left paramere of V. lundbladi); endosoma
without apical accessory sclerite, usually without third
pair of lateral sclerites (but in V. karen with a thin
third pair); in V. kuiterti and V. karen with yellowish
bands on antennal segments 3-4 in both sexes, but
sometimes in heavily pigmented females totally dark.
Distribution (map 4). - Endemic to the Southeast
Asian mainland.
14. Ventidius {Ventidioides) kuiterti Hungerford &
Matsuda
(figs. 130, 135-137, 142, 148, 157, 158, 167, 168, 177,
map 4)
Ventidius {Ventidioides) kuiterti Hungerford & Matsuda,
1960: 333-336 (desc, illustr., key).
Type locality. - Myanmar: Shingbwiyang.
Type material examined. - Pararypes: Myanmar [Bur-
ma]: 46 45, apt., Shingbwiyang, 24.ii.1944, leg. L.C.
Kuitert (semc, jtpc, nhmw); 1 c? apt. Tingkawk, V.1944,
leg. L.C. Kuitert (jtpc).
Other material examined. - Myanmar: ld 49 apt.,
Shingbwiyang, 22.ii.1944, leg. L.C. Kuitert (semc, nhmw);
1 6 1 9 apt., Tingkawk, V.1944, leg. L.C. Kuitert (semc).
Redescription
Dimensions. - Apterous form: length 2.46 (<5),
2.60 (9), width 1.60 (<?), 1.90 (9), width of head
1.10 (<?), 1.20 ($).
Colour (fig. 130). - Whole body prominently dark,
with distinct yellowish marks. Eye blackish. Interocu-
lus blackish, with one large M-shaped yellowish mark
at posterior margin. First antennal segments dark ex-
cept basal 1/6; second segment dark; segment 3 most-
ly yellowish, fourth segment brownish or yellowish at
basal half. Pronotum dark, in some specimens with
two small yellowish marks at the lateroposterior mar-
gin. Mesonotum yellowish with broad dark lateral
stripe which confluent with the dark mark on metan-
otum. Metanotum with two triangular yellowish
marks laterally. Mesopleura with variable colour pat-
Figs. 135-141. Lateral view of apterous males, showing the colour patterns of pleura.
karen; 139, V. sp. from Viet Nam; 140, V. pulai, 141, V. lundbladi.
135-137, Ventidius kuiterti; 138, V.
179
Tijdschrift voor Entomologie, volume mi, 1998
tern; one apterous paratype male with totally yellow-
ish mesopleura (fig. 135); one aprerous paratype male
with a small brownish mark near the anterior margin
of mesopleura (fig. 136); one apterous paratype female
with broad dark stripe running through the length of
mesopleura (fig. 137). Distal margin of metapleura
brownish. External part and lower part of metacetab-
ula blackish (figs. 157, 158). Fore leg dark with basal
Vis yellowish (fig. 148). Middle and hind legs dark.
Tergites 1-3 completely dark, 4 blackish with a trian-
gular yellowish mark in the middle, 5-6 yellowish but
dark laterally, 7 yellowish, 8 dark. In males: lateroter-
gites 1-4 blackish, 5 dark with a small pale mark, 6-7
yellowish; connexiva 1-5 dark, 6-7 yellowish. In fe-
males: laterotergites and connexiva yellowish. Venter
yellowish. Metasternum yellowish.
Pilosity. - Inner surface of antennal segment 1 with
3 subbasal and 1 subapical spines, in male through
basal half of segment 1 to segment 4 with dark fine
hair fringe; segments 2-4 with scattered brownish
short setae (fig. 142). Dorsum, pleura and metacetab-
ula bearing dark pubescence. Venter clothed by gold-
en pubescence, especially on genital segments, the pi-
losity longer and denser. In male at the end of
mesosternal groove with a small tubercle, which is
covered by short dark setae. Long stiff spines scattered
along middle and hind legs.
Figs. 142-147. Dorsal view of male right antennae. - 142,
V. kuiterti; 143, V. karen; 144, V. sp., female, from Viet
Nam; 145, V. pulai, male; 146, same species, female; 147, V.
lundbladi.
Figs. 148-154. Dorsal view of right fore leg. - 148, V.
kuiterti; 149, V. karen; 150, V. sp. female, from Viet Nam;
151, V. pulai, male; 152, V. pulai, female; 153-154, V.
lundbladi; 153, dorsal view; 154, posteroventral view of fore
femur, showing the tooth.
180
Chen & Zettel: Revision ofVentidius
Structural characters. - Interoculus subequal to
width of an eye, 0.40 : 0.40 in male, 0.45 : 0.40 in fe-
male, respectively. Antennae not modified, measure-
ments see table 1. Thorax not bulbous, mesonotum
somewhat swollen, its lateral width 0.90 (6), 1.30
(9). Intersegmental suture between meso- and
matenota obscure. Metanotum somewhat declivent.
Lower part of metacetabula bilobate (figs. 157, 158).
Metasternum not prominent. Fore femur of male
slightly incrassate, halfway its inner surface with a
small tooth (fig. 148), claw rising from 2/5 of seg-
ment 2 of fore tarsus. Anterior margin of abdominal
tergite 1 obscure, anterior margins of tergites 2 and 3
obscure but eventually obliterated medially and pro-
truding forward; abdominal sternite 7 as long as the
preceding abdominal sternites together in both sexes.
Laterotergites broad.
Male terminalia. - Parameres (figs. 167, 168) asym-
metrical, both hooked subbasally, left paramere larger
than the right one, with blunt apex, extending beyond
genital segments. Endosoma (fig. 177): dorsal sclerite
long and recurved proximally, ventral sclerite long,
lateral sclerites straight, hooked distally and broad-
ened proximally, second lateral sclerite long and thin.
Female terminalia. - Abdominal sternite 7 large,
posterior margin smooth, anteriorly concave.
Macropterous form. - Unknown.
Distribution (map 4). - Myanmar: Shingbwayang.
India: Arunachal Pradesh.
Comparative notes. - This species is most similar
to V. karen, for differences see under comparative
notes ofthat species.
15. Ventidius {Ventidioides) karen Lansbury
(figs. 131, 132, 138, 143, 149, 159, 160, 169, 170,
178, 181, 183, map 4)
Ventidius karen Lansbury, 1988: 61-66 (descr., illustr.);
Zettel & Chen, 1996: 152, 181 (list, ree).
Type locality. - Thailand: Nakhon Ratchasima.
Type material examined. - Holotype, â , apt., Thailand:
[Nakhon Ratchasima], Khao Yai National Park, Pha Kuai
Mai Waterfall, 6.xi.l987, leg. C. Deelman (oumc).
Paratype 6 , apt., same data as holotype (oumc).
Other material examined. - Thailand: 3d 99 all apt.,
17lv., Phetchabun Prov., Huai Nam Phang, S. Ban Nam
Nao, 25.xi.1995, leg. H. Zettel; ca. 31 km NW der Abzwei-
gung von der Straße Lom Sak-Chumpae, ca. 5-6 m breiter
Fluß, sehr stief, sehr schlammig (nhmw, ppcc); 5d 3 9 all
apt., Petchabun Prov., Nam Nao N.P., Huai Phrom Laeng,
2-3 m breiter Bach, 24.xi.1995, leg. H. Zettel (22) (nhmw);
75 d, 90 9 all apt., viet nam: : Già Lai-Kontum Prov., 40
km NW An Khe, Buon Luoi, 14°10' N., 108°30' E., 620-
750 m, 28.iii.-12.iv.1995, leg. Pacholatko & Dembicky
(nhmw, nctn, ppcc, bmnh, zrcs).
Figs. 155-156. V. pulai. - 155, dorsal view of fore wing (af-
ter Cheng); 156, dorsal view of hind wing (after Cheng).
163
Figs. 157-166. Male right metacetabula: odd numbers: dor-
sal view; even numbers: dorsolateral view. - 157, 158, Ven-
tidius kuiterti; 159, 160, V. karen; 161, 162. V sp., male,
from Viet Nam; 163, 164, V. pulai; 165, 166, V lundbladi.
181
Tijdschrift voor Entomologie, volume hi, 1998
Figs. 167-176. External view of parameres, odd numbers: right paramere (except 173: internal view), even numbers: left para-
mere (except 174: posterior view of right paramere). - 167, 168, Ventidius kuiterti; 169, 170, V. karen; 171-174, V. pulai;
175, 176, V. lur,
181
180
Figs. 177-180. Lateral view of endosoma sclerites. - 177, Ventidius kuiterti; 178, V. karen; 179, V. pulai; 180, V. lundbladi.
-Fig. 181. V. karen, ventral view of male 8th abdominal segment. - Fig. 182. V. lundbladi, dorsal view of male 7th abdomi-
nal tergite, showing the pilosity.
182
Chen & Zettel: Revision ofVentidius
Redescription
Dimensions. - Apterous form, length 2.66 (a),
3.17 (9), width 1.72 (<?), 2.17 (9), width of head
1.15(d), 1.30(9).
Colour (fig. 1 83). - Whole body predominantly yel-
lowish dorsally, with dark marks. Eyes mainly dark
brown to blackish, with brownish margins. Interocu-
lus dark from dorsal view, with a thick M-shaped yel-
lowish mark at its posterior margin, along inner mar-
gin of eye with a dark stripe. Antennal segments
mainly blackish, but yellowish at basal Vi of segment
1, most part of the segment 3 and middle part of
fourth segment; in female sometimes all antennal seg-
ments blackish except basal Vi of first segment yellow-
ish. Pronotum blackish, lateroposterior angle with a
very small yellowish mark. Mesonotum with broad
lateral dark stripes, which are confluent with the later-
al triangular blackish mark on metanotum. Metan-
otum also with a large triangular dark mark basomedi-
ally which is close to the anterior margin. Mesopleura
yellowish without longitudinal stripe at anterior half
(fig. 138); posterior margin of metapleura dark; upper
external half and lower external half of metacetabula
dark (figs. 159, 160). Fore leg dark, femur with basal
2/5 yellowish (fig. 149). Middle and hind legs dark.
Tergites 1-2 blackish, 3-4 blackish with obscure yel-
lowish mark in the middle, 5-6 dark laterally with a
median yellowish mark, 7 yellowish, with posterior
margin dark, 8 dark. Laterotergites and connexiva 1-5
blackish, 6-7 yellowish in male, predominantly yel-
lowish with dark margins in female. Venter yellowish,
male segment 8 ventrally dark (fig. 132).
Pilosity. — Inner surface of antennal segment 1 with
4 subbasal and 1 subdistal long spine, around seg-
ments 2-4 clothed with usual pubescence, also with
longer setae and spinules. In male from distal half of
segment 1, through all antennal segments with fine
silvery hair fringe. Dorsum and pleura bearing dark
pubescence. Long stiff spines scattered along middle
and hind legs, tibia with shorter stiff spinules, inner
surface of tarsal segment 1 of middle leg with a fringe
of hairs (fig. 149). Basal part of middle and hind
femora with two very long trichobothria-like seta on
dorsal surface respectively, the external one shorter
than the internal one. Ventrally clothed by golden
pubescence, on genital segments the pilosity slightly
longer and denser.
Structural characters. - Interoculus subequal to
width of an eye, 0.39 : 0.36 in male, 0.46 : 0.42 in fe-
male, respectively. Antennae slender, segment 2 short-
er than segment 3 (Measurements see table 1); segment
3 in male not modified at distal half (fig. 143). Prono-
tum not bulbous, mesonotum strongly swollen dorsal-
ly, its lateral width 1.04 (â), 1.36 (9). Mesosternal
groove conspicuous, distally terminated by a small tu-
bercle covered by short brown setae. Intersegmental su-
Fig. 183. Ventidius (Ventidioides)
male, length 2.66 mm.
paratype, apterous
ture between meso- and metanotum faint but visible.
Metanotum somewhat declivent. Lower half of hind
margin of metacetabula bilobate (figs. 159, 160).
Metasternal tubercle not prominent in male. Fore fe-
mur slightly curved, ventral surface with a small tooth
(fig. 149). Anterior margin of abdominal tergite 1 visi-
ble; anterior margins of tergites 2 and 3 faint, obliterat-
ed medially and protruding forward; abdominal stern-
ite 7 as long as the preceding abdominal sternites
together in both sexes. Laterotergites broad.
Male terminalia. - Parameres (figs. 171, 172) asym-
metrical, both hooked at subbasal part, left paramere
larger than right one, its apex twisted and blunt; right
paramere bar-shaped with blunt apex; not extending
beyond genital segments. Endosoma (figs. 178, 181):
dorsal sclerite long and recurved proximally, ventral
sclerite long, lateral sclerite straight, hooked at distal
and broadened at proximal parts, second lateral scle-
rite long and thin, third lateral sclerites curved, apical
sclerite indistinct.
Female terminalia. - Abdominal sternite 7 large,
183
Tijdschrift voor Entomologie, volume i4i, 1998
broadly elevated medially, posterior margin slightly
concave forward.
Macropterous form. - (According to Lansbury,
pers. comm.) 3.60 mm long, maximum width 1.90
mm. colour pattern (fig. 131) pronotum black with
two irregular yellowish blotches. Wings anteriorly
black with fine pubescence and with scattered longer
black hairs. Posterior membranous part of wings dark
brown. The other morphological characters as in
apterous form except the apex of pronotum pointed
(measurements see table 3); length of fore wing from
humeri to apex 2.60.
Distribution (map 4). - Thailand: Nakhon Rat-
chasima, Petchabun; Viet Nam: Gia Lai-Kontum.
Comparative notes. - Ventidius karen shares most
characters with V. kuiterti, especially the short anten-
nal segment 3, the yellowish bands on antennal seg-
ments 3 and 4, and the mesosternal groove ending
with a small tubercle which is covered by dark setae.
Main differences between these species are found in
body length (V. kuiterti smaller than V. karen), in
dorsal colour patterns of meso- and metanotum (in V.
kuiterti darker than in V. karen), in relative lengths of
antennal segment 3 and 4 (segment 4 longer in V.
karen), and in male parameres (see figs. 169, 170).
Furthermore, V. karen is the only species within the
V. kuiterti-group, which has a third pair of endosoma
lateral sclerites (see fig. 178).
Remarks. - One apterous female (figs. 139, 144,
150, 161, 162, 184) was collected from Viet Nam: :
Prenn, 900 m, 15 km S of Da Lat, No. 294,
15.X.1988, leg. Vasarhehj (tmbc). This specimen has
been listed as a Ventidius ( Ventidioides) sp. by Zettel
& Chen (1996: 152, 181). As male specimens are
lacking, it is difficult to decide its specific status. Ac-
cording to its aberrant appearance, we present the de-
scription below:
Dimensions. - apterous female: length 3.30, width
2.35, width of head 1.38.
Colour (fig. 184). - Whole body predominantly
yellowish, with dark marks. Eyes mainly greyish. In-
teroculus yellowish with a broad blackish transversal
mark, along inner margin of eye with a dark stripe.
Antennal segments mainly blackish, but yellowish at
basal lA of the segment 1 , distal half of the segment 3
and middle part of the segment 4 with a yellowish
band respectively. Pronotum blackish, lateroposterior
angle with a small yellowish mark, posterior margin
with two small yellowish marks. Mesonotum yellow-
ish without dark stripes. Metanotum with a triangu-
lar dark mark basomedially. Mesopleura yellowish
with interrupted dark stripe (fig. 139). Anterior exter-
nal half and posterior external angle of metacetabula
dark (figs. 161, 162). Fore leg dark, femur with basal
2/5 yellowish (fig. 150). Middle and hind legs dark.
Tergite 1 black in the middle, and yellowish laterally,
Fig. 184. Ventidius {Ventidioides) sp. from Viet Nam, apter-
ous female, length 3.25 mm.
2-3 blackish with triangular yellowish mark in the
middle, 4 yellowish with dark lateral and anterior
margins, 5-6 and 8 yellowish with dark lateral mar-
gins, 7 completely yellowish. Laterotergites complete-
ly yellowish and connexiva 2-4 blackish, 5-7 yellow-
ish. Venter yellowish.
Pilosity. - Inner surface of antennal segment 1 with
4 subbasal and 1 subapical long spines, around seg-
ments 2-4 clothed with usual pubescence, also with
longer setae, especially on segment 4. Dorsum and
pleura bearing dark pubescence. Long stiff spines
scattered along middle and hind legs. Tibia with
shorter stiff spinules.
Structural characters. - Interoculus broader than
width of an eye, 0.52 : 0.42. Antennae slender, seg-
184
Chen & Zettel: Revision ofVentidius
ment 2 shorter than segment 3 (fig. 144) (measure-
ments see table 1). Pronotum not bulbous, mesono-
tum strongly swollen dorsally, lateral width of
mesothorax 1.40 (9). Intersegmental suture between
meso- and metanotum not visible. Metanotum some-
what declivent. Lower half of hind margin of metac-
etabula bilobate (figs. 161, 162). Fore femur slightly
curved, ventral surface not modified (fig. 150). Ante-
rior margins of abdominal tergite 1-3 not visible.
Female terminalia. - Abdominal sternite 7 large, as
long as the preceding abdominal sternites together,
broadly elevated medially, posterior margin slightly
concave forward.
Comparative notes. - This specimen is similar to
V. karen in most characters. But differences are found
in colour of pronotum, mesonotum, and abdominal
tergites, which are much darker in V. karen. The tho-
rax is much more bulbous than in V. karen. It is diffi-
cult to decide until the male specimen will be avail-
able. Although the specimen represents most likely a
further species of the V. kuiterti-group, it is not im-
possible that it is a very aberrant female of V. karen.
16. Ventidius pulai Cheng
(figs. 133, 140, 145, 146, 151, 152, 155, 156, 163,
164, 171-174, 179, 185, map 4)
Ventidius pulai Cheng, 1965: 153-155, 163 (descr., illustr.,
key); Kovac &C Yang 1989: 285 (ree); Yang & Kovac
1995: 293 (ree).
Type locality. - Malaysia: Johor.
Type material examined. - Holotype c? , apt., allotype 9 ,
apt., and pararypes (30*1$ apt.): Malaysia: Johor, stream
at Gunung Pulai, 500 m, 17.iii.1963, leg. L. Cheng (bmnh,
zrcs).
Other material examined. - Malaysia: 15c?, 25 5 apt.,
3 c?, 2 9 macr., Perak, stream 58 km S. of Grik, CL 2077,
19.viii.1985, J.T. & D.A. Polhemus (jtpc, nhmw);
Malaysia: 1 9 apt., Perak, Kerunai River, 9 km N of Grik at
bridge, CL 2078, 19.viii.1985, J.T. & D.A. Polhemus
(jtpc); lc? 2 9 apt., 2 9 macr., Perak, Ipah Hill, stream nr.
reservoir, 4.iii.l927 (bmnh, nhmw); 1 c? 19 apt., 1 9
macr., Perak, Sungai Batang Padang, nr. for. camp,
13.iii.1927, coll. Dover (bmnh); 3 9 apt., Selangor, Klang
River, near Klang Gates, 21 .viii. 1926, coll. Dover (bmnh);
4cT apt., Selangor, Ampang, waterworks, 5.ÌX.1926, coll.
Dover (bmnh, zmuc); 19 apt., same locality, 30. ii. 1927
(bmnh); 2<5 49 Selangor, Sungai Ampang, 5.ix. 1926, coll.
Dover (bmnh, nhmw, zmuc); 6c? 8 9 apt., same locality,
15.viii.1926, coll. Dover (bmnh, ppcc); 1 6 apt., same lo-
cality, 28.viii.1926, coll. Dover (bmnh); 24c? 40 9 apt., 1 S
1 9 macr., Selangor, Templer Park, CL 2070, 17.viii.1985,
leg. J.T. &c DA. Polhemus (jtpc, nhmw, ppcc); lc?, 3 9
macr., Johor, Gunong Pulai, 26. ii. 1961, leg. C.H. Fernando
(semc); 79 apt., Johor, Endau, Sg. Anak Jasin, 4.iv.l992,
L186, leg. H.K Lua (zrcs, nhmw); 2c? 2 9 apt., same lo-
cality data, leg. K.L. Yeo (zrcs); 5 9 apt., Johor, S Taku,
185
Tijdschrift voor Entomologie, volume i4i, 1998
Fig. 185. Ventidius (Ventidioides) pulai, paratype, apterous
male, length 2.63 mm.
18.vii.1989, K.L. Yeo (zrcs); 19 apt., Johor, S Taku,
15.vi.1989, leg. C.M. Yang et al. (zrcs); 1 9 macr., Negeri
Sembilan, 10 km N Simpang Pertang, unnamed stream,
10.iv.1994.A- 1063, leg. W.D. Shepard (nhmw); lc? 1?
macr., Terengganu, Sg. Terengganu tributary, Sekayu,
16.V.1995, leg. Bruce Tan & Sumita, TG6 (zrcs); 1 9 apt.,
Kelantan, Sungai Hulu Besut, 6 km (W?) after Kampang
Keruah, on road to hydroelectric station, 20.iii.l992, LI 84,
leg. H.K. Lua (zrcs); 20 5 9 apt., 19 macr., Pahang, S.
Kinchin, 19.vii.1989, leg. K.L. Yeo & K.H. Lua (zrcs,
nhmw); 1<? 39 apt., Pahang, S. Kernam, 19.vii.1989, KL.
Yeo (zrcs); 3â 49 apt., 19 macr., Pahang, S. Kinchin,
13.vi.1989, leg. C.M. Yang et al. (zrcs, ppcc); 3â 39 apt.,
2 9 macr., Pahang, S. Seladang, 19.vii.1989, leg. K.L. Yeo
(zrcs, nhmw); 26 3 9 apt., 8 larvae, Prov.?, H.T. Pagden,
BM 1959-236 (bmnh); LAOS: 16 49, (Indo-China), Lu-
ang prabang, Pak Lueng, 5.iü-1920, leg. R.v. de Salvaza,
1920-280 (bmnh, nhmw); 1 9 apt., Xieng Khouang, Sala
Hat Sampong, 17.xi.1917, leg. R.v. de Salvaza (bmnh);
Thailand: 1 6 3 9 apt., Loei, Kaeng Hai, Nam San Khe,
9.iii.l994, A-1035, leg. W.D. Shepard (css, nhmw). \3
19, apt. Chiang Mai, e. 10 km E Samoeng (18°51'N,
98°38'E), river, leg. W.G. Ullrich (Ullrich collection,
nhmw)
Redescription
Dimensions. - Apterous form: length 2.63 (â), j
2.92 (9), width 1.75 (S), 2.00 (9), width of head
1.20 (S), 1.50 (2).
Colour (fig. 18 5). - Whole body prominently dark,
with distinct yellowish marks. Eye blackish. Interocu-
lus blackish, with one large M-shaped yellowish mark
at posterior margin. Antennal segments dark, except
basal 1/5 of segment 1. Prono turn totally dark.
Mesonotum dark with central triangular yellowish
mark which is confluent with the yellowish mark of
metanotum. Metanotum with a triangular dark mark
basomedially and two oblique yellowish stripes lateral-
ly. Mesopleura yellowish, with a small dark mark ante-
riorly; anterior, external margin and lower part of
metacetabula blackish (fig. 140). Metasternum dark.
Fore leg dark with basal 1/5 yellowish (figs. 151, 152).
Middle and hind legs dark. Tergites 1-4 completely
dark, 5-6 blackish with a yellowish mark in the middle,
7 yellowish but dark laterally, 8 dark. Laterotergites 1-
4 blackish, 5 dark with a fine yellowish mark, 6-7 yel-
lowish. Connexiva 1-5 dark, 6-7 yellowish. Sternite 7
dark laterally, segment 8 ventrally dark in male.
Pilosity. — Inner surface of antennal segment 1 with
4 subbasal and 1 subapical spines, in males through
basal half of segment 1 to segment 4 with silvery fine
hair fringe; segments 2-4 with scattered brownish
short setae (figs. 145, 146). Dorsal side, pleura, and
acetabula clothed with dark pubescence. Dark setae
prominent at subdistal part along inner surface of fore
tibia. Venter clothed by golden pubescence, especial-
ly on genital segments the pilosity longer and denser.
Venter yellowish, the setae on mesosternal tubercle
dark brown. Long stiff spines scattered along middle
and hind legs, denser than usual in the genus.
Structural characters. - Interoculus broader than
width of an eye, 0.44 : 0.41 in male, 0.50 : 0.41 in fe-
male. Antennal segment 3 subequal to segment 4
(measurements see table 1) (fig. 145, 146). Thorax
not bulbous, mesonotum somewhat swollen, its later-
al width 1.10 (S), 1.45 (9). Intersegmental suture
between meso- and matanotum obscure. Metanotum
somewhat declivent. Lower part of metacetabula bilo-
bate (figs. 163, 164). Fore femur slightly incrassate,
hallway its inner surface with a small tooth (fig. 151),
claw rising from 2/5 of segment 2 of fore tarsus. Fore
leg of female see fig. 152. Anterior margin of abdom-
inal tergite 1 obscure, anterior margins of tergites 2
and 3 obscure but eventually obliterated medially and
protruding forward; abdominal sternite 7 as long as
the preceding abdominal sternites together in both
sexes. Laterotergites broad.
Male terminalia. - Parameres (figs. 171-174) asym-
metrical, stout, curved at basal part, left paramere dis-
tinctly longer than right one, its apical part curved up-
wards, with blunt apex; apical part of right paramere
186
Chen & Zettel: Revision ofVentidius
Fig. 186. Ventidius (Ventidioides) lundbladi, apterous male,
length 2.37 mm.
straight and pointed upwards, with blunt apex; the left
paramere extending beyond genital segments. Endo-
soma (fig. 179): dorsal sclerite long and recurved prox-
imally, ventral sclerite long, lateral sclerites straight,
broadened at two ends, second lateral sclerites thin
and weak.
Female terminalia. - Abdominal sternite 7 large,
posterior margin smooth, concave forwards.
Macropterous form. - Maximum width 1.66 mm.
colour pattern: Pronotum black with two regular yel-
lowish marks (fig. 133); wings black anteriorly with
fine pubescence and with scattered longer black hairs;
posterior membranous part of wings dark brown
(figs. 155, 156). Other morphological characters as in
apterous form except the apex of pronotum pointed
(measurements see table 3).
Distribution (map 4). - Thailand: Loei; Laos; Ma-
laysia: Perak, Johor, Selangor, Pahang, Kelantan,
Negeri Sembilan, Terengganu.
Comparative notes. - Typical member of the V.
kuiterti-group, and easy to distinguish from V. kuiter-
ti and V. karen by subequal and uniformly brown an-
tennal segments 3 and 4 in the male, and a less devel-
oped blotch of dark setae on mesosternum. The left
paramere is apically twisted (fig. 174) similar as in V.
karen (fig. 170), but less curved. The shape of para-
meres, a usually darker colouration, and a less devel-
oped pilosity of segment 8 of males are the main dif-
ferences to V. lundbladi (see under comparative notes
ofthat species).
17. Ventidius (Ventidioides) lundbladi Miyamoto
(figs. 134, 141, 147, 153, 154, 165, 166, 174-176,
180, 182, 186, map 4)
Ventidius lundbladi Miyamoto, 1967: 245-247 (descr., illustr.).
Type locality. - Thailand: Khao Chong.
Material examined. -Thailand: 8 S 1 1 9 apt., S. Banna,
Nakhon, 108 m, 5-10.V.1958, leg. T.C. Maa, No. 415
(bimc, jtpc, nhmw, ppcc).
Redescription
Dimensions. - Apterous form. Length 2.37 (<?),
2.60 (9), width 1.57 (<?), 1.88 (9), width of head
1.04(d), 1.11 (9).
Colour (fig. 186). - Whole body prominently
dark, with distinct yellowish marks. Eye blackish. In-
teroculus pale, with one large heart-shaped dark
mark, the dark mark along inner margin of eye reach-
ing posterior margin of vertex, and connected with
the heart-shaped one. Antennal segments dark, ex-
cept basal 2/5 of segment 1. Pronotum usually com-
pletely dark, in some specimens with two small yel-
lowish marks at the lateroposterior margin of
pronotum. Mesonotum with lateral dark mark.
Metanotum with triangular dark mark. Mesopleura
yellowish (fig. 141); external angle of metacetabula
dark (figs. 165, 166). Under strong light, along the
intersegmental suture between metanotum and ter-
gite 1 , the pubescence reflects with a shining greenish
metallic colour. Fore leg dark with basal lA yellowish
(figs. 153, 154). Middle and hind legs dark. Tergites
1-4 completely dark, 5 yellowish with lateral dark
mark, 6-7 yellowish, 8 dark. Laterotergites 1-4 dark,
5 dark basally, 6-7 yellowish. Connexiva 1-5 dark, 6-
7 yellowish. Venter yellowish.
Pilosity. - Trichobothria prominent (fig. 134) In-
ner surface of antennal segment 1 with 3 subbasal, and
1 subapical long spines; in males through basal half of
segment 1 to apical half of segment 4 with silvery fine
hair fringe; segments 2-4 with scattered dark short se-
tae (fig. 147). Dorsum and pleura clothed with dark
pubescence on yellowish marks, and golden pubes-
cence on dark parts. Venter clothed by golden pubes-
cence, especially on genital segments, the pilosity very
long and dense. Long stiff spines scattered along mid-
dle and hind legs. Basal part of middle femur with a
very long trichobothria-like seta on dorsal surface.
Structural characters. - Interoculus subequal to
width of an eye, 0.40 : 0.32 in male, 0.40 : 0.35 in fe-
male. Antennal segment 3 subequal to segment 4
187
Tijdschrift voor Entomologie, volume mi, 1998
Fig. 187. Ventidius (Ventidioides) xyele, paratype, apterous
male, length 2.95 mm.
(measurements see table 1) (fig. 147). Thorax not bul-
bous, mesonotum somewhat swollen, its lateral width
1.00 (c?), 1.33 (9). Intersegmental suture between
meso- and metanotum distinct. Metanotum some-
what declivent. Lower part of metacetabula bilobate
(figs. 165, 166). Metasternal tubercle prominent in
male. Fore femur slender and with a small tooth ante-
rio-ventrally, claw rising from 2/5 of segment 2 of fore
tarsus. Anterior margin of abdominal tergite 1 dis-
tinct; anterior margins of tergites 2 and 3 obliterated
medially and protruding forward; abdominal sternite
7 as long as the preceding abdominal sternites togeth-
er in both sexes. Laterotergites broad.
Male terminalia. - Parameres (fig. 175, 176) asym-
metrical, both hook-shaped, left paramere longer
than right one, its apex sharp and pointed forward,
distal part of right paramere straight upwards, with
blunt apex; both parameres extending beyond genital
segments. Endosoma (fig. 180): dorsal sclerite long
and curved proximally, ventral sclerite long, lateral
sclerite straight, broadened at distal part, second lat-
eral sclerites thin and weak, apical sclerite indistinct.
Female terminalia. - Abdominal sternite 7 large,
posterior margin smooth, concave forwards.
Macropterous form. - Unknown.
Distribution (map 4). - Thailand: S. Banna, Khao
Chong.
Comparative notes. - Although the types were not
available for our study, there is no doubt about this
species which is characterized by the shape of para-
meres figured by Miyamoto (1967: figs. 73-74). It is
most similar to V. pulai, but easily distinguishable by
the shape of parameres: left paramere (fig. 174) long
and slender in distal part, not plate-like and apically
not twisted as in V. pulai; right paramere as in fig.
175. The dense developed long dark pilosity on seg-
ment 8 of male (fig. 182) richer than in V. pulai; Both
sexes are also different in the thoracic colour pattern,
which is usually much darker in V. pulai.
The Ventidius xiphibion-group
Diagnosis. - Prominent dark colouration; male fore
femur without sharp tooth; most species with modi-
fied male fore leg, two species with a thickening on
fore tibia, other species with tuft of black setae on fore
trochanter, one species ( V. heissi sp. n.) without both
of these characters; mesosternum with a patch of dark
setae along median mesosternal groove; parameres
protruding beyond genital segments, larger than in
other Ventidius species, more or less asymmetrical; en-
dosoma with distinct third pair of lateral sclerites, in
most species with distinct apical accessory sclerite.
Notes. - This group includes five species, two from
Sulawesi and three from Borneo. The Sulawesian
species are characterized by a tuft of black setae on male
fore trochanter and by asymmetrical parameres, which
are strongly enlarged basally. Two of the Bornean
species are characterized by a modified male fore tibia,
which is thickened and with dense short pilosity in
middle of its length, by a distinct swelling of male fore
femur, and by less asymmetrical, hooked parameres.
Ventidius heissi sp. n. is regarded as related to these
species by the general characters of male genitalia.
Distribution (map 5). - Borneo and Indonesia:Su-
lawesi.
18. Ventidius ( Ventidioides) xiphibion Chen & Nieser
(figs. 188, 193, 198, 203, 204, 213, 216, 219, 220,
229, 230, map 5)
Ventidius xiphibion Chen & Nieser, 1992: 156-157 (descr.,
illustr.).
188
Chen & Zettel: Revision ofVentidius
Figs. 188-192. Lateral view of apterous male, showing the
colour pattern of pleura. - 188, Ventidius xiphibion; 189, V.
xyele, 190, V. kurtokalami; 191, V. nieseri; 192, V. heissi.
Figs. 193-197. Dorsal view of male right antennae. — 193,
Ventidius xiphibion; 194, V. xyele, 195, V. kurtokalami; 196,
V. nieseri; 197, V. heissi.
Type locality. — Indonesia: Sulawesi.
Type material examined. - Holotype 3 , apt., allotype ?
apt., Indonesia: Sulawesi Tenggara Prov., small stream 8
km E of Sungai Sampara along road Kendari-Wawotobi,
N8911, 22.ii.1989, leg. N. Nieser (rmnh); Paratypes: 41 ó\
64 9 , apt., 1 â macr., same locality data as holotype (rmnh,
nctn, ppcc, nhmw); 35 S 39$, apt., 1$ macr., Sulawesi
Tenggara Prov., P. Buton, stream just N of Bau-bau,
N8925, 8.iii.l989, leg. N. Nieser (rmnh, nctn, nhmw);
40c? 175 apt., 1 5 macr., Sulawesi Tenggara Prov., stream
about 15 km E of Bau-bau, N8942, 10.iii.1989, leg. N.
Nieser (rmnh, nctn).
Other material examined. — 1 3 1 5 apt., Sulawesi Ten-
gali Prov., Ampana, 31.1.1995, leg. Seyfert & Graindl (49)
(nhmw).
Locality notes. - Small to medium sized streams with tur-
bid water due to suspended loam. The species tends to ag-
gregate in places with little current.
Redescription
Dimensions. - Apterous form. Length 2.90 (â),
3.15 (9), width 2.00 (c?), 2.26 (9), width of head
1.28 (cT), 1.28(9).
Colour. - Whole body prominently dark, with dis-
tinct yellowish marks. Eye brownish. Interoculus
189
Tijdschrift voor Entomologie, volume mi, 1998
Figs. 1 98-202. Dorsal view of right fore leg.
202, V. heissi.
198, Ventidius xiphibion; 199, V. xyele, 200, V. kurtokalami; 201, V. nieseri;
213/
Figs. 203-212. Metacetabula, odd numbers: dorsal view,,
even numbers: dorsolateral view. - 203, 204, Ventidius
xiphibion; 205, 206, V. xyele, 207, 208, V. kurtokalami; 209,
210, V. nieseri; 211, 212, V. heissi. —Fig. 213. V. xyele, ven-
tral view of fore trochanter, showing the tuft of dark setae.
blackish, with one large M-shaped yellowish mark at
posterior margin. Antennal segments dark, except basal
2/5 of segment 1 . Pronotum dark in male, in some
specimens with a small central yellowish mark; in fe-
male dark, with a central and two small yellowish
marks at the lateroposterior margin, in some specimens
the posterior margin with faint yellowish marks.
Mesonotum yellowish with broad lateral dark stripe
which is confluent with the dark mark of metanotum.
Metanotum blackish with two oblique yellowish
marks. Mesopleura yellowish, with a small black mark
near anterior margin in male (fig. 188), with a brown-
ish stripe running through its length in female, but in
some specimens the dark marks are obscure. External
half of metacetabula blackish (figs. 203, 204). Fore leg
dark with basal Vi yellowish (fig. 198). Middle and
hind legs dark. Tergites 1-4 completely dark, 5 black-
ish with a triangular yellowish mark in the middle, 6-7
yellowish with lateral and posterior margin dark, 8
dark. Laterotergites 1-2 blackish, 3-7 yellowish. Con-
nexiva 1-4 dark, 5-7 yellowish. In female the coloura-
tion of abdomen paler than in males. Venter yellowish.
Pilosity. - Inner surface of antennal segment 1
with 6-8 subbasal and 1 subapical spines, through
basal half of segment 1 to segment 4 with fine silvery
hair fringe; segments 2-4 with scattered brownish
short setae (fig. 193). Dorsum, pleura and metacetab-
ula bearing dark pubescence. Ventral surface of fore
trochanter with a tuft of dark setae apically (fig. 213).
190
Chen & Zettel: Revision ofVentidius
Figs 214-218. -214, Ventid-
his kurtokalami, dorsal view of
macropterous female (ap-
pendages removed); 215, V.
nieseri, ventral view of body,
showing the patch of dark
mesosternal setae; 216-218,
dorsal view of macropterous
pronotum. - 216, V.
xiphibion; 217, V. nieseri;
218, V.xyele.
Figs. 219-228, External view
of parameres, odd numbers:
left paramere, even numbers:
right paramere. - 219, 220,
Ventidius xiphibion; 221, 222,
V. xyele, 223, 224, V. kur-
tokalami; 225, 226, V. nieseri;
227, 228, V. heissi.
191
Tijdschrift voor Entomologie, volume i4i, 1998
Figs. 229-234. - 229, Ventid-
ius xiphibion, dorsal view of
endosoma sclerites. - 230-
234. Lateral view of endoso-
ma sclerites. — 230, V.
xiphibion; 231, V. xyele, 232,
V. kurtokalami; 233, V.
nieseri; 234, V. heissi
229
233
Dark setae along inner surface of fore tibia not modi-
fied (fig. 198). Venter clothed by golden pubescence,
especially on genital segments the pilosity longer and
denser. On half way of mesosternal groove with a
blotch of dark setae which is displayed in two longi-
tudinal lines. Stiff spines scattered along middle and
hind legs.
Structural characters. - Interoculus broader than
width of an eye, 0.47 : 0.40 in male, 0.47 : 0.42 in fe-
male. Antennae not modified, measurements see table
1 (fig. 193). Thorax not bulbous, mesonotum some-
what swollen, its lateral width 1.17 (c?), 1.35 (9). In-
tersegmental suture between meso- and metanotum
distinct. Metanotum somewhat declivent. Lower part
of metacetabula bilobate (figs. 203, 204). Metasternal
tubercle not prominent. Fore femur incrassate, taper-
ing towards distal part (fig. 198), claw rising from Vs
of segment 2 of fore tarsus. Anterior margin of ab-
dominal tergite 1 obscure but visible, anterior margins
of tergites 2 and 3 obscure but eventually obliterated
medially and protruding forward; abdominal sternite
7 as long as the preceding abdominal sternites togeth-
er in both sexes. Laterotergites broad.
Male terminalia. - Parameres (figs. 219, 220) asym-
metrical, both hooked, left paramere longer and
broader than right one, knife-shaped, tapering distal-
ly, with sharp apex. Right paramere slender and with
blunt apex. Endosoma (figs. 229, 230): dorsal sclerite
long and recurved proximally, ventral sclerites long
and weak, lateral sclerites straight, broadened at two
ends, second lateral sclerites thin and weak, third lat-
eral sclerites distinct and curved; apical accessory scle-
rite distinct.
Female terminalia. - Abdominal sternite 7 large,
posterior margin smooth, concave forwards.
Macropterous form. - Maximum width 1.75 mm.
Colour pattern: pronotum (fig. 216) black with two
regular yellowish blotches. Wings anteriorly black
with fine pubescence and with scattered longer black
hairs. Posterior membranous part of wings dark
brown. The other morphological characters as apter-
ous form except the apex of pronotum pointed (mea-
surements see table 3); length of fore wings from
humeri to apex 2.40.
Distribution (map 5). - Indonesia: Sulawesi Teng-
gara Prov., Sulawesi Tengah Prov.
192
Chen & Zettel: Revision ofVentidius
Map 5. Ventidius {Ventidioides)
xipbibion-group.
• xiphibion
O xyele
■ kurtokalami
□ nieseri
•k heissi
Comparative notes. — Most characters of this
species are similar to that of V. xyele, especially the less
modified male fore leg, the shape of left and right
parameres which are hooked upward close to a strong-
ly enlarged base and very asymmetrical (left paramere
much broader than right one). These characters unite
both Sulawesian species and separate them from the
Bornean species of the V. xiphibion-group. Differences
between V. xiphibion and V. xyele are found in the ex-
tensive yellow pattern (much darker in V. xyele) and
different shapes of male parameres: parameres of V.
xyele ha.ve very slender and sharp apex (fig. 221, 222),
in V. xiphibion the apex is not so sharply constricted
(fig. 219, 220). The endosoma of V. xiphibion has a
distinct apical accessory sclerite which is less promi-
nent in V. xyele, but clearly present in the Bornean
species of the subgenus.
19. Ventidius {Ventidioides) xyele Chen & Nieser
(figs. 187, 189, 194, 199, 205, 206, 218, 221, 222,
231, map 5)
Ventidius xyele Chen & Nieser, 1992: 157-158 (descr. & il-
luso.).
Type locality. - Indonesia: Sulawesi.
Type material examined. - Holotype c?, apt., allotype 9
apt., Indonesia: Sulawesi Utara Prov., Dumoga Bone Na-
tional Park, R. Toraut, Maze, 16.xi.1985, leg. G. Zimmer-
mann (rmnh). Paratypes: 3c? 99, apt., same locality data as
holotype (rmnh, zmac, nctn). 1 o* 1 9 apt., Dumoga Bone
National Park, Base camp, 16.x. 1986; 19, Dumoga Bone
River, downstream, of bridge, 22.X.1985; 8c? 99, apt.,
Tumpah River, Beach, 19-1985; 7c? 79 , apt., Tumpah Riv-
er, Staustufe (low barrier), 23.X.1985; 3c? apt., 2c? 1 9 macr.,
Malibagu Z. 8.xi.l985; 4 c? 5 9 , apt., Südküste, Strasse, Bach
(S. coast, road, rivulet), 18.xi.1985. (deposited separetely in
ZCWA, nctn, bpuh, jtpc, mbbj, ppcc, zmac, zmuc).
Other material examined. - Indonesia: lc? 10 9 apt.,
Sulawesi Utara Prov., P. Sangihe, Naha, Sungai Laine at last
bridge upstream, N9456, 27.vi.1994 (nctn); 12c? 99 apt.,
1 9 macr., Sulawesi Utara Prov., P. Sangihe, D. Simuang
(dekat di Malahu), Sungai Simuang, N9457, 28.vi.1994,
leg. N. Nieser, mountain stream water clear, hyacine, boul-
ders, stones, stretches with sand (nctn); 20 <? 15 9 apt., Su-
lawesi Utara Prov., P. Sangihe, Desa Laine, Sungai Laine
(different from N9454/56 which is on the other side of the
island); pothole as water fall, 12.xi.1994, N9463, leg. N.
Nieser (nctn); 5 c? 129 apt., Sulawesi Utara Prov., P.
Sangihe, small stream at Kampung Lapango-Hakadele (near
esa Sawaeng), 18.xi.1994, N9474, leg. N. Nieser (nctn);
9c? 8 9 apt., Sulawesi Utara Prov., P. Sangihe, Desa Laine,
Sungai Laine, near the road, 18.xi.1994, N9476, leg. N.
Nieser, muddy pebbles between boulders, depth about 1 m,
quiet flow of water, except for some shallow banks with rif-
fles (nctn); 9c? 69 apt., Sulawesi Utara Prov., P. Sangihe,
Sungai Miulu, 20.xi.1994, N9480, leg. N. Nieser (nctn);
27c? 39 9 apt., Sulawesi Utara Prov., P. Sangihe, desa Utau-
rano, Sungai Apanukang, pothole, I4.xi.1994, N9465A,
193
Tijdschrift voor Entomologie, volume i4i, 1998
leg. N. Nieser (nctn); 2 c? 4 9 , apt., Sulawesi Selatan Prov.,
E side Lake Matana, Kg. Salura: Nightjar Camp, 520 m,
13.X.1993, open water and among mangrove vegetation,
2°32'15'S. 121028WE., leg. J.P. & M.J. Duffels (zmac);
2c? 5 9 , apt., same locality data as above, 450 m, 20.x. 1993,
narrow tributary to Lake Matana, 2°32'S., 121°28'E., J.P.
& M.J. Duffels (zmac); 1 cT 2?', apt., SW Sulawesi Selatan
Prov., Onang, Sg. Parabaya, 58 km N of Majene,
19.xi.1993, strongly disturbed rain forest, leg. J.P.& M.J.
Duffels (zmac); AS 79 apt., 1 c? macr., Sulawesi Utara
Prov., cave spring and stream at Komangaan, NW Kota-
mobagu, CL 2120, l4.ix.1985, leg. D.A. Polhemus (jtpc,
nhmw); Ilo* 379 apt., 19 macr., Sulawesi Utara Prov.,
Tondano River tributary, S of Airmadidi, CL 2127,
20.ix.1985, leg. J.T. & D.A. Polhemus (jtpc, nhmw); 49c?
42 9 apt., lc? macr., Sulawesi Utara Prov., tributary to
Tumpah River, CL 2101, 4.ÌX.1985, leg. J.T. & D.A. Pol-
hemus (jtpc, nhmw, ppcc); 5 c? 49 apt., Sulawesi Utara
Prov., Pononontuna River at Tapakulintang, 200 m, CL
2121, 15.ix.1985, leg. J.T. & D.A. Polhemus (jtpc,
nhmw); 33 c? 38 9 apt., Sulawesi Utara Prov., Tumpah Riv-
er, Dumoga-Bone Nat. Park, 0°34'N, 123°54'E, CL 2100,
4.ÌX.1985, 222 m, leg. J.T. & D.A. Polhemus (jtpc,
nhmw); 18c? 28 9 apt, 1 9 macr., Sulawesi Utara Province,
stream near Manembonembo, E of Manado, CL 2128,
20.ix.1985, leg. J.T. & D.A. Polhemus (jtpc, nhmw); 8c?
169 apt., 19 macr., Sulawesi Selatan Prov., Pattunuang
River, 7 km SW of Bantimurung, 0-100 m, CL 2165,
13.X.1985, leg. J.T. & D.A. Polhemus (jtpc, nhmw); 2c?
5 9 apt., Sulawesi Selatan Province, Marana River, nr. Cam-
ba, 50 km E of Maros, 450 m, CL 2167, 14.x. 1985, leg. J.T.
6 D.A. Polhemus (jtpc, nhmw).
Redescription
Dimensions. - Apterous form: Length 2.95 (c?),
3.10 (9), width 2.06 (cT), 2.26 (9), width of head
1.18(d), 1.21 (9).
Colour (fig. 187). -Whole body prominently dark,
with distinct yellowish marks. Eye blackish. Interocu-
lus blackish, with one large M-shaped yellowish mark
at posterior margin. Antennal segments dark, except
basal Va of segment 1 . Pronotum dark in male, in fe-
male lateroposterior margin with a small yellowish
mark. Mesonotum yellowish with broad dark lateral
stripes which are confluent with the dark mark of
metanotum. Metanotum blackish with two oblique
yellowish marks. Mesopleura yellowish, with a longi-
tudinal dark stripe in male (fig. 189), in female this
dark stripe less developed. External half of metacetab-
ula blackish (figs. 205, 206). Fore leg dark with basal
% (c?) (fig. 199) or Yi (9) of femur yellowish. Middle
and hind legs dark. Tergites 1-4 completely dark, 5-7
blackish with yellowish mark in the middle. Lateroter-
gites 1-3 blackish, 4 blackish with yellowish mark, 5-7
yellowish. Connexiva 1-4 blackish, 5-7 yellowish.
Venter yellowish. Segment 8 darkened ventrolaterally.
Pilosity. - Inner surface of antennal segment 1 with
5-6 subbasal and 1 subapical spines, through basal
half of segment 1 to segment 4 with fine silvery hair
fringe; segments 2-4 with scattered brownish short se-
tae (fig. 194). Dorsum and pleura and metacetabula
bearing dark pubescence. Ventral surface of fore
trochanter with a tuft of dark setae apically (fig. 213).
Dark setae along inner surface of fore tibia not modi-
fied (fig. 199). Venter clothed by golden pubescence,
especially on genital segments, the pilosity longer and
denser. Halfway the mesosternal groove with a dark
setae blotch which is displayed in two longitudinal
lines. Stiff spines scattered along middle and hind
legs, denser than in most other species.
Structural characters. - Interoculus broader than
width of an eye, 0.49 : 0.41 in male, 0.50 : 0.44 in fe-
male. Antennae not modified, measurements see table
1 (fig. 194). Thorax not bulbous, mesonotum some-
what swollen, its lateral width 1.12 (c?), 1.27 (9). In-
tersegmental suture between meso- and metanotum
obscure. Metanotum somewhat declivent. Lower part
of metacetabula bilobate (figs. 205, 206). Metasternal
tubercle not prominent. Fore femur of male incras-
sate, tapering towards distal part (fig. 199), Claw ris-
ing from basal Vi of segment 2 of fore tarsus. Anterior
margin of abdominal tergite 1 obscure but visible, an-
terior margins of tergites 2 and 3 obscure but eventu-
ally obliterated medially and protruding forward; ab-
dominal sternite 7 as long as the preceding abdominal
sternites together in both sexes. Laterotergites broad.
Male terminalia. — Parameres (figs. 221, 222) asym-
metrical, both hooked, left paramere longer and
broader than right one, knife-shaped, tapering distal-
ly, with sharp apex. Right paramere slender and with
sharp apex. Endosoma (fig. 231): dorsal sclerite long
and recurved proximally, ventral sclerites long and
weak, lateral sclerites straight, broadened at both ends,
second lateral sclerites thin and weak, third lateral scle-
rite long and curved, apical accessory sclerite small,
not visible from lateral view.
Female terminalia. - Abdominal sternite 7 large,
posterior margin smooth, concave forwards.
Macropterous form. - Maximum width 1.70 mm.
colour pattern: pronotum (fig. 218) black with two
regular yellowish blotches. Wings anteriorly black
with fine pubescence and with scattered longer black
hairs. Posterior membranous part of wings dark
brown. Other characters as apterous form except the
apex of pronotum pointed (measurements see table
3); length of fore wings from humeri to apex 2.30.
Distribution (map 5). - Indonesia: Sulawesi Utara
Prov., Sulawesi Selatan Prov.
Comparative notes. - Most closely related to V.
xipbibion; for difference see the comparative notes
under that species.
20. Ventidius ( Ventidioides) kurtokalami Chen &
Nieser
(figs. 190, 195, 200, 214, 223, 224, 232, 235, map 5)
194
Chen & Zettel: Revision ofVentidius
Ventidius {Ventidioides) kurtokalami Chen & Nieser,
158-159 (descrip. &: illustr.).
Type locality. - Malaysia: Sabah, Danum Valley.
1992:
Type material examined. - Holotype <? , apt., allotype 9 ,
apt., E. Malaysia: Sabah, Danum Valley, 70 km W Lahad
Datu, 4 km S main trail W5 nr. Sungai Segama, 150 m,
middle sized stream and waterfall, 3.xii. 1 989, sample Sab.
54, M.J. & J. P. Duffels (zmac). Paratype: lc? apt., Sabah,
Danum Valley, 70 km W Lahad Datu, main trail W 12, 180
m, narrow creek, 2.XÜ.1989, sample Sab. 52, M.J. & J. P.
Duffels (nctn).
Other material examined. - Malaysia: 15 c? 22$ apt.,
1 e? macr., Sabah, Danum Valley, Palum Tambun, 2.-
13.2.1997, leg. H. Zettel (2) and leg. Zettel et al. (different
project numbers) (umsm, spcm, nhmw); 3<? 5 9 Sabah,
Danum Valley, Sapat Kalisan, 12.2.1997, leg. H. Zettel
(15) (umsm, nhmw); 8c? 11 9 apt. Sabah, Danum Valley,
Waterfall, 3.2.1997, leg. H. Zettel (5) (umsm, nhmw); 5 c?
69 apt., Sabah, Danum Valley, Water Pool, 1 1.2.1997, leg.
H. Zettel (13) (umsm, nhmw); 1 1 c? 3 9 apt., Sabah, North
Borneo (SE), Forest Camp, 19 km N of Kalabakan,
12. xi. 1962, leg. Y. Hirashima (bimc, nhmw); 5c? 109 apt.,
from same locality, 13.xi.1962 (bimc, ppcc); 5c? 2 9 apt.,
Sabah, North Borneo, Tawau, Quoin Hill, Forest Camp,
1,3-5 km WSW of Cocoa Res. Sta., 9.-20.viii.l962, leg. Y.
Hirashima (bimc, nhmw); 2 9 apt., Sabah, North Borneo,
Tawau, Quoin Hill, Cocoa Res. Sta., 26.ix.1962, leg. Y. Hi-
rashima (bimc); 1 c? 2 9 apt., Sabah, British North Borneo,
Tawau, Quoin Hill, 26.-29.vii. 1962 (bimc); 1 â 1 9 apt.,
Sabah, Samlang River, 7 km S of Ranau, CL 2026,
3.viii.l985, leg. J.T. & D.A. Polhemus (jtpc); 5 c? 79 apt.,
1 c? 1 9 macr., Sabah, 40 km NE of Kota Belud, CL 2032,
5.viii.l985, leg. J.T. & D.A. Polhemus (jtpc, nhmw); 1 9
apt., Sabah, trib. to Moyog River nr. km 12 on Keningau
Hwy., CL 2039, 6.viii.l985, leg. J.T. & D.A. Polhemus
I (jtpc); 5 c? 3 9 apt., Sabah, Maliau Basin, fast flow stream,
MB9a and MB10, 16.5.1996, leg. T.B. Lim (zrcs, nhmw).
Redescription
Dimensions. - Apterous form. Length 2.90 (<3),
3.45 (9), width 2.00 (<?), 2.60 (9), width of head
1.25 (6), 1.30 (2).
Colour (fig. 235). - Whole body prominently yel-
lowish, with distinct dark marks. Eye brown to black-
ish. Interoculus yellowish, with one large heart-
shaped dark mark at middle. Antennal segments
dark, except basal 1/6 of segment 1, and the colour of
segment 4 paler. Pronotum dark, only with two small
yellowish marks at the lateroposterior margin.
Mesonotum yellowish with broad dark lateral stripes
which are confluent with the dark mark of metan-
otum. Metanotum with a triangular dark mark medi-
ally and two transverse blackish bands laterally.
Mesopleura yellowish; upper and lower external an-
gles of metacetabula brownish (fig. 190). Fore leg
dark with basal Vi of femur yellowish. Middle and
hind legs dark. Tergites 1-3 completely dark, 4 black-
ish with an obscure triangular yellowish mark in the
middle, 5-7 yellowish but dark laterally, 8 yellowish
with posterior margin dark. Laterotergites yellowish,
Fig. 235- Ventidius {Ventidioides) kurtokalami, paratype,
apterous male, length 2.90 mm.
3-4 more or less darkened. Connexiva 1-5 dark, 6-7
yellowish. Venter yellowish.
Pilosiry. - Inner surface of antennal segment 1 with
2 subbasal and one subapical spines, through basal half
of segment 1 to segment 3 with dark fine hair fringe;
segments 2-4 with scattered brownish short setae (fig.
195). Dorsum and pleura clothed with dark pubes-
cence on yellowish marks, and golden pubescence on
dark part. Dark setae along both inner and external
surfaces of fore tibia very prominent in male, which
give the inner surface of fore tibia a wavy outline (fig.
200). Venter clothed by golden pubescence, especially
on genital segments the pilosity longer and denser.
Along mesosternal groove with a row of short dark
bristles. Long stiff spines scattered along middle and
hind legs.
Structural characters. - Interoculus subequal to
width of an eye, 0.44 and 0.42 in male, 0.50 and 0.46
195
Tijdschrift voor Entomologie, volume i4i, 1998
in female. Antennae not modified, measurements see
table 1 (fig. 195). Thorax not bulbous, mesonotum
somewhat swollen, its lateral width 1.10 (6), 1.60
(9). Intersegmental suture between meso- and
metanotum obscure. Metanotum somewhat de-
clivent. Lower part of metacetabula bilobate (figs.
207, 208). In the centre of male mesonotum with a
small patch of minute black bristles which are not on
a tubercle. Metasternal tubercle prominent in male.
Fore femur slightly incrassate, with a blunt hump
(fig. 200). Because of the prominent pilosity the fore
tibia of male modified along inner and external mar-
gin, inner surface with a hump halfway its length,
which is covered by erect pubescence, claw rising
from 2/5 of segment 2 of fore tarsus. Anterior margin
of abdominal tergite 1 obscure, anterior margins of
tergites 2 and 3 obscure but eventually obliterated
medially and protruding forward; abdominal sternite
7 as long as the preceding abdominal sternites togeth-
er in both sexes. Laterotergites broad.
Male terminalia. - Parameres (figs. 223, 224) asym-
metrical, left paramere longer than right one, hooked,
apical half curved upwards, with blunt apex. Endoso-
ma (fig. 232): dorsal sclerite long and recurved proxi-
mally, ventral sclerite long, lateral sclerite straight,
broadened and hooked at two ends, third lateral sclerite
recurved backwards, apical accessory sclerite distinct.
Female terminalia. - Abdominal sternite 7 large,
posterior margin smooth, concave forwards.
Macropterous form. - (fig. 214). Its maximum
width 1.85 mm. Colour pattern: Pronotum black
with two regular yellowish blotches separated by a thin
brownish line. Wings anteriorly black with fine pubes-
cence and with scattered longer black hairs. Posterior
membranous part of wings dark brown. Other charac-
ters as in apterous form except the apex of pronotum
pointed (measurements see table 3); length of fore
wings from humeri to apex 2.15.
Distribution (map 5). - East Malaysia: Sabah.
Comparative notes. - This species is very similar to
V. nieseri sp. n., both sharing the modified fore leg in
males. For differences see comparative notes of V.
nieseri sp. n.
21. Ventidius {Ventidioides) nieseri sp. n.
(figs. 191, 196, 201, 209, 210, 215, 217, 225, 226,
233, 236, map 5)
Type locality. - Brunei; Malaysia: Sarawak.
Type material. - Holotype 6, apt., allotype 5,
apt., Brunei: Kuala Belalong Field Research Centre,
I6.iv.1993, N.9345, leg. N. Nieser (rmnh). -
Paratypes: 46 2 9 , all apt., same locality data as holo-
type (rmnh, nhmw, nctn, ppcc); 3Ó* 1 9 apt., 1 9
macr., dars, Temburong, Belalong FR Centre, small
stream behind FRC, l6.-17.iv.1993, N9345A, leg.;
N. Nieser (nctn); 36 1 9 apt., 1 ó* 1 9 macr., Tern- !
burong, Kuala Belalong Field Research Centre, Main
river, N9344, I6.iv.1993, leg. N. Nieser, quiet edge;
downstream of boat & jetty (nctn); 26 apt., Tem-|
brong, Kuala Belalong Field Research Centre, sungai
Baki, well sized tribitary to main river, 17.iv.1993,
leg. N. Nieser, N9364 (nctn, nhmw); 1 9 apt., 96 \
4 9 macr., Temburong, Belalong Field Res. Centre,
Sungai Belalong, 60 m, 2-8.V.1995, leg. E. Heiss
(zcwa, ppcc, nctn); 1 6 1 9 apt., 1 S 3 9 macr., Sg.
Belalong, Kuala Belalong Field Studies Centre,
I6.vi.1995, leg. S.L. Goh, GSL9504 (zrcs, nhmw);
29 apt., 20* macr., same locality, l4.vi.1996,
GSL9501 (zrcs); Malaysia: 3o* apt., Sarawak, Mulu
N. P., right tributary to Tutoh river, Long Iman,
4.ÌU.1993, leg. H. Zettel (14) (nhmw). 10c? 149,
apt, Sarawak, Batang Ai N.P., Engkari River, E of
Bandar Sri Amman, 19-20.ii.1993, leg. H. Zettel (7)
(nhmw). 26 , apt., Sarawak, Kapit Dist., Merirai V.,
30-300 m, l-6.viii.1958, leg. T. Maa (semc); 26 2 9
apt., Sarawak, Kapit District, Merirai V., 30-300 m,
L-6.viii.1958, secondary forest, leg. T.C. Maa
(bimc); Indonesia: 36 29 apt., 36 49 macr., C.
Borneo (Kalimantan), Sg. Birang, leg. Mjöberg 1925,
Coll. Dr. D. Mac Gillavry (zmac, nhmw, ppcc).
Etymology. - This species is named after our friend
and colleague Dr. Nico Nieser for his devoted excel-
lent work on aquatic and semiaquatic Heteroptera.
Also for his generous offering of numerous interesting
specimens from Sulawesi and Brunei, among them a
large sample of this species.
Locality notes. — Small shaded river, bottom main-
ly pebbles, at margin and foot of waterfall.
Description
Dimensions. - Apterous form. Length 2.78 (â),
2.68 (9), width 1.81 (c?), 1.93 (9), width of head
122(6), 1.25(9).
Colour (fig. 236). - Whole body prominently
dark, with distinct yellowish marks. Eye blackish. In-
teroculus blackish, with one large M-shaped yellow-
ish mark at posterior margin. Antennal segments
dark, except basal 1 IG of segment 1 . Pronotum dark,
only with two small yellowish marks at the lateropos-
terior margin of pronotum. Mesonotum yellowish
with lateral dark stripe which is confluent with the
dark mark of metanotum, but not reaching its anteri-
or margin. Metanotum with a triangular dark mark
medially and two transverse blackish bands laterally.
Mesopleura yellowish. External margin and lower
part of metacetabula blackish (fig. 191). Fore leg dark
with basal 1/5 of femur yellowish (fig. 201). Middle
and hind legs dark. Tergites 1-3 completely dark, 4
196
Chen & Zettel: Revision ofVentidius
Fig. 236. Ventidius ( Ventidioides) nieseri, holotype, apterous
male, length 2.78 mm.
blackish with a triangular yellowish mark in the mid-
dle, 5 yellowish but dark laterally, 6-7 yellowish with
posterior margin dark, 8 dark. Laterotergites 1-4
blackish, 5-7 yellowish. Connexiva 1-5 dark, 6-7 yel-
lowish. Venter yellowish.
Pilosity. - Inner surface of antennal segment 1 with
4 subbasal and 1 subapical long spines, through basal
half of segment 1 to segment 3 with dark fine hair
fringe; segments 2-4 with scattered brownish short se-
tae (fig. 196). Dorsum and pleura bearing golden pu-
bescence on yellowish marks, metacetabula with dark
pubescence. Dark setae along inner surface of fore tib-
ia very prominent, which gives the inner surface of
fore tibia a wavy outline (fig. 201). Venter clothed by
golden pubescence, especially on genital segments the
pilosity longer and denser. Along medium line of
mesosternum with a dark stripe composed of short se-
tae, setae pointed upwards and gradually tapering
onto the anterior margin of mesosternum (fig. 215).
Long stiff spines scattered along middle and hind legs.
Structural characters. - Interoculus subequal to
width of an eye, 0.46 : 0.42 in male, 0.49 : 0.46 in fe-
male. Antennae not modified, but segment 1 distinct-
ly curved towards ventral side, measurements see table
1 (fig. 196). Thorax not bulbous, mesonotum some-
what swollen, its lateral width 1.43 (â), 1.53 (9). In-
tersegmental suture between meso- and metanotum
obscure. Metanotum somewhat declivent. Lower part
of metacetabula bilobate (figs. 209, 210). Metasternal
tubercle hidden under mesosternum. Fore femur of
male incrassate, halfway its inner surface with a blunt
swelling (fig. 201), claw rising from 2/5 of segment 2
of fore tarsus. Anterior margin of abdominal tergite 1
obscure, anterior margins of tergites 2 and 3 obscure
but eventually obliterated medially and protruding
forward; abdominal sternite 7 as long as the preceding
sternites together in both sexes. Laterotergites broad.
Male terminalia. - Parameres (fig. 225, 226) asym-
metrical, left paramere longer than right one, both
hooked, apex of right paramere blunt, apex of left
paramere sharper than right one; paramere extending
beyond genital segments. Endosoma (fig. 233): dorsal
sclerite long and recurved proximally, ventral sclerite
long, lateral sclerites straight, broadened at distal ends,
second lateral sclerites thin and weak, third lateral scle-
rite curved; apical accessory sclerite distinct.
Female terminalia. - Abdominal sternite 7 large,
posterior margin smooth, concave forwards.
Macropterous form. - Its maximum width 1.90
mm. Colour pattern: Pronotum (fig. 217) black with
two regular yellowish blotches. Wings anteriorly black
with fine pubescence and with scattered longer black
hairs. Posterior membranous part of wings dark
brown. Other characters as in apterous form except the
apex of pronotum pointed (measurements see table 3);
length of fore wings from humeri to apex 2.65.
Distribution (map 5). - Brunei; East Malaysia: Sa-
rawak; Indonesia: Kalimantan.
Comparative notes. - Ventidius nieseri sp. n. is
closely related to V. kurtokalami. Differences are small
and mainly found in males: The thickening in the
middle of fore tibia is poorly developed in V. nieseri sip.
n. and mainly indicated by a short erect pilosity; the
fore femur is more curved basally in V. nieseri sp. n.;
the setae along mesosternal groove are more conspicu-
ous in V. nieseri sp. n.; both parameres are slightly
more curved and distally somewhat more slender in V.
nieseri sp. n.; and the endosoma of V. nieseri sp. n. has
a larger apical accessory sclerite; the species are usually
also separable in colour, which is brighter in both sex-
es in V. kurtokalami., but specific variability of both
species is slightly overlapping.
22. Ventidius {Ventidioides) heissi sp. n.
(figs. 192, 197,202,211,212,218,227,228,234,
237, map 5)
Type locality. - Malaysia: Sarawak.
197
Tijdschrift voor Entomologie, volume i4i, 1998
Fig. 237. Ventidius ( Ventidioides) beissi, holotype, apterous
male, length 2.73 mm.
Type material. - Holotype 6 , apt, Malaysia: Bor-
neo / Sarawak, Kapit Dist., Merirai Village, 30-300
m, 1-6. VIII. 1958, secondary forest, No. M.B. 164,
leg. T.C. Maa (bimc). - Paratypes, 46 , apt., same lo-
cality data as holotype (bimc, nhmw, ppcc).
Etymology. - This species is named after our friend
and colleague Prof. Mag. Dr. Ernst Heiss (Innsbruck),
specialist in Oriental Aradidae, who provided us with
interesting specimens from Brunei.
Description
Dimensions. - Apterous male. Length 2.73 (a),
width 1.85 (<J), width of head 1.12 (S).
Colour (fig. 237). - Whole body prominently
dark, with distinct yellowish marks. Eye blackish. In-
teroculus pale, with one large round dark mark. An-
tennal segments 1-3 dark except basal 2/5 of segment
1, segment 4 paler. Pronotum completely dark.
Mesonotum with lateral dark mark. Metanotum with
triangular dark mark. Mesopleura with a thick dark
stripe (fig. 192; external angle of metacetabula dark
(figs. 211, 212). Under strong light, the pubescence
reflects with a shining greenish metallic colour. Fore
leg dark with basal Vi yellowish (fig. 202). Middle
and hind legs dark. Tergites 1 completely dark, 2-5
yellowish but dark laterally, 6 mainly yellowish, 7 yel-
lowish, posterior margin dark, 8 dark. Laterotergites
1 dark, 2-5 dark laterally, 6-7 yellowish. Connexiva
1-5 dark, 6-7 yellowish. Venter yellowish.
Pilosity. - Inner surface of antennal segment 1 with
3 subbasal, and 1 subapical long spines; through basal
half of segment 1 to apical half of segment 4 with fine
hair fringe; segments 2-4 with scattered dark short se-
tae (fig. 197). Dorsum and pleura bearing dark pubes-
cence on yellowish marks, and golden pubescence on
dark parts. Venter clothed by golden pubescence, es-
pecially on genital segments, the pilosity very long and
dense. Halfway the mesosternal groove with a blotch
of dark setae which is displayed in two longitudinal
lines, not as obvious as in other species of this group.
Dark setae and long spines scattered along middle and
hind legs.
Structural characters. - Interoculus subequal to
width of an eye, 0.42 : 0.39 in male. Antennae not
modified, measurements see table 1 (fig. 197). Tho-
rax not bulbous, mesonotum somewhat swollen, its
lateral width 1.28 (a). Intersegmental suture be-
tween meso- and metanotum obscure. Metanotum
somewhat declivent. Lower part of metacetabula bilo-
bate (figs. 211,212). Metasternal tubercle not promi-
nent in male. Fore femur slender, inner surface of fore
tibia not modified, claw rising from 2/5 of segment 2
of fore tarsus. Anterior margin of abdominal tergite 1
indistinct; anterior margins of tergites 2 and 3 oblit-
erated medially and protruding forward; abdominal
sternite 7 as long as the preceding abdominal sternites
together. Laterotergites broad.
Male terminalia. - Parameres (figs. 227, 228) asym-
metrical, both hook-shaped, left paramere longer than
right one, its apex more or less sharp; distal part of
right paramere less pointed, with blunt apex; both
parameres extending beyond genital segments. Endo-
soma (fig. 234): dorsal sclerite long and curved proxi-
mally; ventral sclerites long; first lateral sclerites
straight, broadened at distal parts; second lateral scle-
rites thin and weak; third lateral sclerite oblique, thin
and clear; apical accessory sclerite distinct.
Macropterous form and female. - Unknown.
Distribution (map 5). - East Malaysia: Sarawak.
Comparative notes. - This new species is similar to
V. xiphibion and V. xyele in having a row of setae
along mesosternal groove (although this is less promi-
nent than in any other member of the V. xiphibion-
group) and the lack of a thickening or tubercle on
fore femur and fore tibia in male. The lack of a tuft of
setae on fore trochanter, the long antennal segment 3,
and the general shape of the parameres are characters
which clearly distinguish V. heissi sp. n. from both
198
Chen & Zettel: Revision ofVentidius
Figs 238-244. - 238, Ventidius {Ventidiopsis) imadatei Miyamoto, dorsal view of apterous female (appendages removed),
length 2.90 mm. - 239, 241, 243, dorsal view of metacetabula; 240, 242, 244, dorsolateral view of metacetabula. - 239, V.
imadatei, female; 240, male; 241, 242, V. yangae, male; 243, 244, female.
these Sulawesian species and show a closer relation-
ship to the following two Bornean species, V. kur-
tokalami and V. nieseri sp. n. Males of V. heissi sp. n.
are easy to distinguish from both of these species by
the lack of special structures on fore femur and fore
tibia (fig. 202), the less prominently developed setae
along mesosternal groove, and the broad shape of the
parameres: left paramere apically curved, right para-
mere apically blunt (figs. 227, 228).
Ventidius ( Ventidiopsis) Miyamoto, 1967, stat. n.
Ventidiopsis Miyamoto, 1967: 247. Type species by mono-
type Ventidiopsis imadatei Miyamoto, 1967. Andersen
1982: 393 (classification).
Diagnosis. - Relatively dark-coloured, pronotum
black; metacetabula bilobate; sexes very different: fe-
males strongly modified: thorax dorso-ventrally
swollen, metacetabula with a mediad, hairy processus;
metacoxa with mediad processus; abdomen shortened,
most tergites fused, sternite 7 strongly modified, with
two elongate lateral projections and two caudally
pointed medial lobes. Males similar to the subgenus
Ventidioides, but antennae much stouter, especially
the long segment 1 .
Distribution (map 6). - Endemic to Borneo.
Key to apterous specimens of the subgenus
Ventidiopsis
1. Female with tuft of black bristles on posterior
margin of mesonotum, with acutely pointed me-
dial processus of metacetabula (fig. 240), with
longer processus of metacoxa (fig. 238), and with
long lateral projections of sternite 7 (fig. 263);
male with longer left paramere (fig. 258) and
broader distal part of right paramere (fig. 257)
(Borneo: Brunei, Sarawak) V. imadatei
Female without tuft of black bristles on posterior
margin of mesonotum, with rectangular medial
processus of metacetabula (figs. 241, 242), with
shorter processus of metacoxa (fig. 265), and with
short, appressed lateral projections of sternite 7
(fig. 264); male with shorter left paramere (fig.
260) and more slender distal part of right paramere
(fig. 259) (Borneo: Sabah) V. yangae sp. n.
The Ventidius imadatei-group
Notes. - Only two, very closely related species are
known, which are so far the only representatives of
the subgenus. For diagnostic characters see diagnosis
of the subgenus. Macropterous morphs are unknown.
199
Tijdschrift voor Entomologie, volume mi, 1998
f' 245
246
248
Figs. 245-248. Lateral view of body, showing the colour pattern of pleura. - 245, Ventidius imadatei, male; 246, female; 247,
V. yangae, male; 248, female.
23. Ventidius {Ventidiopsis) imadatei Miyamoto
comb. n.
(figs. 238-240, 245, 246, 249, 250, 253, 254, 257,
258,261,263, map 6)
Ventidiopsis imadatei Miyamoto, 1967: 247-250 (gen. n.,
sp. n., descr., illustr.)
Type locality. - Brunei.
Material examined. — 3c? 1 2 apt., Brunei: Temburong,
Kuala Belalong Field Res. Centre, main river, N9344,
16.iv.1993, leg. N. Nieser, quiet edge downstream of boat
& jetty (nctn, ppcc); 2c? apt., Temburong, Kuala Belalong
Field Research Centre, Sungai Bald, well sized tribitary to
main river, 17.iv.1993, leg. N. Nieser, N9364 (rmnh,
nctn). 5 c? apt., Temburong, Belalong Field Res. Centre,
Sungai Belalong, 60 m, 2-8.V.1995, leg. E. Heiss (hcia,
zcwa, ppcc); 4<? apt., Sg. Belalong, Kuala Belalong Field
Studies Centre, 16.vi.1995, leg. S.L. Goh, GSL9504 (zrcs);
3c? 49 apt., same locality, I4.vi.1995, leg. S.L. Goh,
GSL9501 (zrcs, nhmw); 1 cT 1 9 apt., Sarawak, Kapit Dis-
trict, Merirai V., 30-300 m, l.-6.viii.l958, secondary forest,
leg. T.C. Maa (bimc), 9 c?, 69, apt., Sarawak, Mulu NP, 3-
5.3.1993, leg. H. Zettel (14), (nhmw, ppcc); 1$, apt.,
Sarawak, Lambir Hills NP, 25 km S Miri, 24-25.2.1993, leg
H. Zettel (9), (nhmw); Malaysia: le? apt., Sabah, North
Borneo (SE), Forest Camp, 19 km N of Kalabakan,
12.xi.1962, leg. Hirashima (bimc); 1 9 apt., Sabah, Maliau
Basin, fast flow stream, MB9a, 16.5.1996, leg. T.B. Lim
(umsm); 23, 19 apt., Sabah, Tibow Estate, slow flow
stream, MB42, 25.5.1996, leg. T.B. Lim (zrcs, umsm).
Material not examined (checked by John T. Polhemus,
pers. communication). - Malaysia: 1 9 apt., Sarawak, Sam-
eran River, 2 km W of Tubeh, CL 2047, 19.viii.1985, leg.
J.T. & D.A. Polhemus (jtpc); 2c? 49 apt., Sabah, tributary
to Moyog Rivet, near km 12 on Keningau Highway, CL
2039, 6.viii.l985, leg. J.T. & D.A. Polhemus (jtpc).
Description
Dimensions. - Apterous form. Length 2.73 (<5),
2.90 (9), width 1.83 (cî), 1.80 (9), width of head
1.15(5), 1.23(9).
Colour (fig. 238). - Whole body prominendy dark,
with distinct yellowish marks. Eye blackish. Interocu-
lus blackish, with one large M-shaped yellowish mark
at posterior margin. Antennal segments dark, except
base of segment 1 . Pronotum dark, only with two small
yellowish marks at the lateroposterior margin of prono-
tum. Mesonotum in male yellowish with lateral dark
stripe which is confluent with the dark mark of metan-
otum and reaching its anterior margin. Mesonotum in
female broadly yellowish with lateral dark stripe which
is confluent with the dark mark of metacetabula.
Metanotum of male with a triangular dark mark medi-
ally and two transverse blackish bands laterally. Metan-
otum in female blackish at anterolateral corner, inner
side of lateral longitudinal elevations. Mesopleura yel-
lowish with a dark stripe running through its length
(figs. 245, 246); upper and lower external angles of
metacetabula blackish (figs. 239, 240). Fore leg dark
with basal Va-Vì of fore femur yellowish (figs. 253,
254). Middle and hind leg dark. In males, tergites 1-3
completely dark, tergite 4 blackish with a triangular
yellowish mark in the middle, tergite 5 yellowish but
dark laterally, 6-7 yellowish with posterior margin
dark, tergite 8 dark. Laterotergites 1-5 blackish, lateral
plates 6 -7 yellowish. Connexiva 1 -5 dark, 6-7 yellow-
ish. In females, tergites 1-3 black with broad yellowish
lateral marks, tergites 4-8 yellowish with blackish later-
al margins. Venter yellowish in both sexes.
Pilosity. - Inner surface of male first antennal seg-
200
Chen & Zettel: Revision ofVentidius
Figs. 249-252.
Dorsal view of male right an-
tennae; 253-256. Dorsal view
of right fore leg. - 249, 253,
Ventidius imadatei, female;
250, 254, male; 252, 255, V.
yangae, female; 251, 256,
male.
Figs. 257-264.
257-260, external view of
right (257, 259) and left (258,
260) parameres; 261, 262,
lateral view of endosoma scle-
rites; 263, 264. Vllth sternite
of female. - 257, 258, 261,
263, Ventidius imadatei; 259,
260, 262, 264, V. yangae.
201
Tijdschrift voor Entomologie, volume hi, 199s
ment with 2-3 subbasal spines, but without a subapi-
cal spine as usual case in other Ventidius species (fig.
249). In females, along inner surface of first antennal
segment with 5-6 long spines (fig. 250); in males,
through basal half of segment 1 to segment 3 with
dark fine hair fringe and segments 2-4 with scattered
brownish short setae. Dorsum and pleura bearing
dark pubescence on yellowish marks, and greyish pu-
bescence on dark parts. Inner surface of fore tibia
with suberect pilosity (figs. 253, 254). Females with
two pairs of tufts consisting of long stiff black setae on
posterior margin of mesonotum and on inner projec-
tion of metacetabula. Metacoxa of female on inner
projection with numerous long black hairs. Venter
clothed by dark pubescence. Long stiff spines scat-
tered along middle and hind legs.
Structural characters. - Interoculus slightly broader
than width of an eye, 0.46 : 0.41 in male, 0.43 : 0.41
in female. Male antenna thick (fig. 249), segment 1
distinctly curved towards ventral side in male, mea-
surements see table 1. Thorax bulbous in female,
mesonotum strongly swollen, which makes the female
appear thick from lateral view, its lateral width 1.07
(S), 1.40 (?). Intersegmental suture between meso-
and metanotum obscure. Metanotum declivent.
Metacetabula bilobate in lower part, in female with a
sharp, internally directed processus (fig. 240). Fore fe-
mur slender in both sexes, claws rising from Vi of seg-
ment 2 of fore tarsus. In males anterior margin of ab-
dominal tergite 1 visible, anterior margins of tergites 2
and 3 obliterated medially and protruding forward. In
females abdomen shortened, sutures between tergites
1 -7 obscure or only laterally visible; Abdominal stern-
ite 7 in male shorter than the preceding abdominal
sternites together, in female longer and modified. Lat-
erotergites broad in male, narrow in female.
Male terminalia. - Parameres (fig. 257, 258) asym-
metrical, left paramere twice as long as right one, ex-
tending beyond genital segments, its apical part
curved upwards, with blunt, laterad curved apex;
right paramere short, with pointed apex. Endosoma
(fig. 261): dorsal sclerite long and recurved proximal-
ly, and broadened and homogeneous in dorsodistal
half, ventral sclerites short, lateral sclerites straight,
broadened at basal end, second lateral sclerites thin
and weak. Third lateral sclerite curved distally and
broadened at proximal half.
One male from Tibow Estate with an elongate,
sharply ending right paramere seems to be an individ-
ual aberration because a second male from the same
locality has a normally shaped paramere.
Female terminalia. - Apex of abdomen in dorsal
202
Chen & Zettel: Revision ofVentidius
view with distinct, mediae! curved lateral projections
of sternite 7; genital segments in dorsal view con-
cealed under preceding abdominal segments. In ven-
tral view abdominal sternite 7 large, hiding the gono-
coxae completely, with two slender, strongly
developed lateral projections and two large triangular,
sharply pointed medial lobes (fig. 263).
Macropterous form. - Unknown.
Distribution (map 6). - Brunei; East Malaysia:
Sarawak, Sabah.
Comparative notes. - Closely related to the follow-
ing species, for differences see key and comparative
notes of V. yangae sp. n.
24. Ventidius ( Ventidiopsis) yangae sp. n.
(figs. 241-244, 247, 248, 251, 252, 255, 256, 259,
260,262,264-266, map 6)
Type locality. - Malaysia: Sabah.
Type material. - Holotype 9 , apt. and allotype 6 ,
apt., Sabah, Danum Valley, Palum Tambun, 7.-
12.2.1997, leg. Zettel et al. (P90) (umsm). -
Paratypes: le?, 1 $ apt., same locality data (nhmw); 3
6 apt., Sabah, Danum Valley, Segama River, 'Beach',
4.2.1997, leg. H. Zettel (10) (umsm, nhmw); 6â 5 9
apt., Sabah, Danum Valley, Palum Tambun, 7.-
12.2.1997, leg. Zettel et al. (different project num-
bers) (umsm, spcm, nhmw); 1 9 apt., Sabah, Danum
Valley, Segama River above Field Centre, 2.2.1997,
leg. H. Zettel (1) (nhmw); 2d 39 apt., Sabah,
Danum Valley, Sapat Kalisan, 12.2.1997, leg. H.
Zettel (15) (nhmw).
Etymology. - Dedicated to Dr. Yang Chang Man
(zrcs) for her continuous help in making type speci-
mens and other specimens available for this study and
for her various contributions to the knowledge of
Malesian water bugs.
Description
Dimensions. - Apterous form. Length 2.82 (cî),
3.00 (9), width 1.65 (<?), 1.82 (9), width of head
1.20 (<J), 1.20(9).
Colour (figs. 265, 266). - Whole body dark, with
distinct yellowish marks. Eye blackish. Interoculus
blackish, with one large M-shaped yellowish mark at
posterior margin. Antennal segments dark, except base
of segment 1 . Pronotum dark, only with two small
yellowish marks at the lateroposterior margin of
pronotum. Mesonotum in male yellowish with lateral
dark stripe which is confluent with the dark mark of
metanotum and reaching its anterior margin.
Mesonotum in female broadly yellowish with lateral
dark stripe which is confluent with the dark mark of
metacetabula. Metanotum of male with a triangular
Fig. 265. Ventidius ( Ventidioides) yangae, holotype, apterous
female, length 2.90 mm.
dark mark medially and two transverse blackish bands
laterally. Metanotum in female blackish at anterolater-
al corner, inner side of lateral longitudinal elevations
with an elongate dark mark medially, which is conflu-
ent laterally with abdominal dark marks. Female
mesopleura yellowish with a dark stripe running
through its length (fig. 247), male mesopleura totally
yellowish or with thin brownish longitudinal stripe
(fig. 248); upper and lower external angles of metac-
etabula blackish (figs. 241-244). Fore leg dark with
basal Vi of fore femur yellowish (figs. 255, 256). Mid-
dle and hind leg dark. In males, tergites 1-3 complete-
ly dark, tergite 4 blackish with a triangular yellowish
mark in the middle, tergite 5 yellowish but dark later-
ally, 6-7 yellowish with posterior margin dark, tergite
8 dark. Laterotergites 1-5 blackish, lateral plates 6-7
yellowish. Connexiva 1-5 dark, 6-7 yellowish. In fe-
males, tergites 1 yellowish with lateral dark marks, 2-7
yellowish, tergite 8 yellowish with dark patches anteri-
orly. Venter yellowish in both sexes.
Pilosity. - Inner surface of male antennal segment
1 with 2-3 subbasal spines, but without a subapical
spine as usual case in other Ventidius species (fig.
203
Tijdschrift voor Entomologie, volume i4i, 1998
Fig. 266. Ventidius ( Ventidioides) yangae, allotype, apterous
male, length 2.73 mm.
252). In females, along inner surface of antennal seg-
ment 1 with 4-5 long spines (fig. 251); in males,
through basal half of segment 1 to segment 3 with
dark fine hair fringe and segments 2-4 with scattered
brownish short setae. Dorsum and pleura bearing
dark pubescence. Inner surface of fore tibia with
suberect piloshy (figs. 255, 256). Females with one
pair of tufts consisting of long stiff black setae on in-
ner projection of metacetabula (fig. 242). Metacoxa
of female on inner projection with numerous long
black hairs. Venter clothed by dark pubescence. Long
stiff spines scattered along middle and hind legs.
Structural characters. - Interoculus slightly broader
than width of an eye, 0.42 : 0.40 in male, 0.50 : 0.46
in female. Male antenna thick (fig. 252), segment 1
distinctly curved towards ventral side in male, mea-
surements see table 1. Thorax bulbous in female,
mesonotum strongly swollen, which makes the female
appear thick from lateral view, its lateral width 1.05
((?), 1.35 ($). Intersegmental suture between meso-
and metanotum obscure. Metanotum declivent, espe-
cially in females. Metacetabula bilobate in lower part,
in female with a rectangular internally directed proces-
sus (figs. 241-244). Fore femur slender in both sexes,
claws rising from Vi of segment 2 of fore tarsus. In
males anterior margin of abdominal tergite 1 visible,
anterior margins of tergites 2 and 3 obliterated medi-
ally and protruding forward. In females abdomen
shortened, sutures between tergites 1-7 obscure or
only laterally visible; Abdominal sternite 7 in male
shorter than the preceding abdominal sternites togeth-
er, in female longer and modified. Laterotergites
broad in male, narrow in female.
Male terminalia. - Parameres (fig. 259, 260) asym-
metrical, left paramere one and a half times as long as
right one, extending beyond genital segments, its api-
cal part curved upwards, with blunt, laterad curved
apex; right paramere short, with pointed apex. Endo-
soma (fig. 262): dorsal sclerite long and recurved
proximally, ventral sclerites short, lateral sclerites
straight, broadened proximally, second lateral scle-
rites thin and weak, which are covered by the third
lateral sclerite, the third lateral sclerite hooked distal-
ly and broadened at proximal half.
Female terminalia. - Apex of abdomen in dorsal
view without distinct projections of sternite 7; genital
segments in dorsal view concealed under preceding
abdominal segments. In ventral view abdominal ster-
nite 7 (fig. 264) large, hiding the gonocoxae anterior-
ly, with two short angular lateral projections which
are appressed to tergite 7, and with two large triangu-
lar, pointed medial lobes, distance between the two
lobes relatively broader comparing with V. (Ventid-
iopsis) imadatai.
Macropterous form. - Unknown.
Distribution (map 6). - East Malaysia: Sabah.
Comparative notes. — Closely related to V. ima-
datei. Main differences are presented in the key. In
addition, the medial lobes of the female sternite 7 are
of different shapes (see figs. 259-264). Differences in
the endosoma are found in the distally hooked third
lateral sclerite of V. yangae sp. n., which is less curved
in V. imadatei.
References
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Gerromorpha). Phylogeny, Adaptation, Biogeography
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Andersen, N. M., 1990. Genus and taxonomy of water
striders, genus Aquarius Schellenberg (Insecta, Hemi-
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Bergroth, E., 1911. On some recently described Hemiptera,
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Chen, P. P. & N. Nieser, 1992. Gerridae, mainly from Su-
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schrift voor Entomologie 135: 145-162.
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Cheng, L., 1966. Three new species of Esakia Lundblad
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Esaki, T., 1928. New or little known Gerridae. I. Ceylonese
species. - Annals and Magazine of Natural History 10:
505-513.
Esaki, T., 1929. New or little known Gerridae. II. Indian
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412-419.
Esaki, T., 1930. New or little known Gerridae from the
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ptera: Gerridae) from India. - Oriental Insects 15: 97-102.
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Hungerford, H. B. & R. Matsuda, 1960. Concerning the
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Received: 20 October 1997
Revised version accepted: 8 June 1998
205
Tijdschrift voor Entomologie, volume i4i, 1998
Table 1. Lengths of antenna! segments in Ventidius species (in mm)
I
II
III
VI
V. aquarius
Ô
1.78
1.21
0.65
0.58
9
1.57
0.80
0.66
0.57
V. harrìsoni
Ô
1.50
0.94
0.75
0.57
9
1.27
0.61
0.64
0.52
V. longitarsus
3
2.00
1.32
0.70
0.88
9
1.80
0.79
0.65
0.79
V. malayensis
â
1.95
1.37
0.65
0.78
9
1.45
0.79
0.58
0.73
V. Usingen
S
1.65
1.32
0.79
0.61
9
1.43
0.82
0.72
0.58
V. bungerfordi
â
1.10
0.67
0.49
0.44
9
1.00
0.42
0.54
0.42
V. polhemorum
â
1.07
0.58
0.49
0.40
9
0.90
0.43
0.51
0.40
V. werneri
6
1.05
0.50
0.40
0.41
9
0.90
0.40
0.47
0.37
V. pilosus
â
1.15
0.53
0.32
0.39
9
1.01
0.43
0.41
0.41
V. henryi
â
0.93
0.41
0.42
0.35
9
0.88
0.34
0.45
0.35
V. chinai
â
1.15
0.56
0.41
0.36
9
1.05
0.44
0.49
0.42
V. modulatus
â
1.20
0.61
0.48
0.39
V. kuiterti
s
1.21
0.39
0.42
0.45
9
1.12
0.32
0.48
0.50
V. karen
â
1.40
0.32
0.35
0.48
9
1.14
0.31
0.43
0.47
Ksp.
9
1.12
0.36
0.50
0.52
V. pulai
â
1.17
0.42
0.48
0.49
9
1.13
0.40
0.58
0.53
V. lundbladi
3
1.18
0.37
0.41
0.40
9
0.96
0.36
0.51
0.44
V. xiphibion
3
1.29
0.47
0.38
0.50
9
1.25
0.48
0.50
0.51
V. xyele
â
1.26
0.50
0.41
0.52
9
1.30
0.54
0.54
0.51
V. kurtokalami
3
1.22
0.57
0.55
0.56
9
1.30
0.50
0.70
0.60
V. nieserì
â
1.30
0.51
0.53
0.50
9
1.29
0.50
0.70
0.60
V. heissi
3
1.15
0.47
0.51
0.53
V. imadatei
â
1.50
0.77
0.57
0.51
9
1.20
0.55
0.60
0.48
V. yangae
â
1.50
0.70
0.52
0.45
1.00
0.96
0.08
0.28
Esakia ventidioidesS
1.05
0.35
0.33
0.28
9
1.10
0.30
0.33
0.30
206
Table 2. Lengths of leg segments in Ventidius species.
Chen & Zettel: Revision of Ventidius
male (c> )
female
(?)
species
femur
tibia
tarsi
femur
tibia
tarsi
V. aquarius
fore leg
1.43
1.30
0.39
1.38
1.26
0.41
mid leg
3.95
2.48
1.45
3.75
2.25
1.40
hind leg
4.18
2.03
0.56
4.00
2.15
0.59
V. usingeri
fore leg
1.60
1.35
0.46
1.46
1.18
0.42
mid leg
4.15
2.51
1.60
3.77
2.40
1.48
hind leg
4.45
1.90
0.62
3.76
2.01
0.68
V. harrisoni
fore leg
1.42
1.15
0.39
1.18
1.05
0.42
mid leg
3.47
2.04
1.20
3.00
1.70
1.07
hind leg
3.50
1.76
0.53
3.03
1.54
0.56
V. malayensis
fore leg
1.77
1.45
0.60
1.50
1.30
0.60
mid leg
4.50
2.75
1.65
4.07
2.46
1.58
hind leg
4.82
2.50
0.70
4.17
2.36
0.75
V. longitarsus sp
. n. (Da
Lat)
fore leg
1.80
1.60
0.77
1.66
1.36
0.64
mid leg
4.90
3.00
1.90
4.45
2.70
1.60
hind leg
4.90
2.65
0.70
4.30
2.56
0.80
V. longitarsus sp
. n. (Gia
Lai-Kontum
)
fore leg
1.53
1.55
0.77
1.51
1.40
0.62
mid leg
4.53
2.88
1.70
4.16
2.60
1.70
hind leg
4.71
2.57
0.92
4.00
4.00
0.88
V. hungerfordi
fore leg
1.11
0.93
0.33
1.10
0.92
0.32
Mid leg
2.75
1.50
1.00
3.00
1.50
1.10
hind leg
3.10
0.70
0.35
3.00
1.30
0.36
V. polhemorum
sp. n.
fore leg
1.05
0.90
0.30
1.05
0.90
0.30
mid leg
2.67
1.28
1.00
2.77
1.43
1.06
hind leg
2.57
1.12
0.40
2.68
1.25
0.45
V. werneri
fore leg
1.03
0.86
0.30
0.95
0.93
0.28
mid leg
2.66
1.42
0.95
2.68
1.41
0.98
hind leg
2.68
1.11
0.38
2.61
1.11
0.35
V. pilosus sp. n.
fore leg
1.08
0.96
0.35
1.13
1.05
0.38
mid leg
3.06
1.78
1.30
3.33
1.90
1.35
hind leg
3.46
1.78
0.58
3.66
1.65
0.65
V. henryi
fore leg
1.06
0.95
0.31
1.05
0.84
0.31
mid leg
3.03
1.76
1.07
3.03
1.80
1.07
hind leg
3.38
1.30
0.41
3.40
1.23
0.41
V. modulatus
fore leg
1.10
0.96
0.35
1.11
0.96
0.34
mid leg
3.00
1.75
1.11
3.26
1.80
1.17
hind leg
3.19
1.37
0.42
3.40
1.50
0.50
V. kuiterti
fore leg
1.06
0.91
0.32
1.10
1.00
0.32
mid leg
3.00
1.16
1.09
3.20
1.75
1.00
hind leg
3.25
1.05
0.45
3.50
0.85
>>
V. karen
fore leg
1.12
0.90
0.30
1.20
0.99
0.38
mid leg
3.00
1.50
1.00
3.25
1.66
1.12
hind leg
3.18
1.17
0.40
3.55
1.42
0.50
V. sp. ( V. karen
> from Viet Nam)
fore leg
pp
pp
pp
1.30
1.04
0.38
mid leg
pp
>>
pp
3.48
1.90
1.22
hind leg
pp
>>
^
3.58
1.58
0.54
continued on
page 208
207
Tijdschrift voor Entomologie, volume i4i, 1998
V. pulai
fore leg
1.11
0.90
0.33
1.11
0.94
0.33
mid leg
3.01
1.61
1.02
3.20
1.75
1.05
hind leg
3.21
1.28
0.45
3.51
1.38
0.51
V. lundbladi
fore leg
1.10
0.86
0.28
1.08
0.90
0.31
mid leg
2.54
1.51
1.05
3.00
1.60
0.94
hind leg
2.76
1.20
0.52
2.97
1.28
0.43
V. xiphibion
fore leg
1.10
0.90
0.35
1.20
0.90
0.30
mid leg
3.20
1.90
1.10
3.30
1.90
1.10
hind leg
3.50
1.50
0.40
3.70
1.60
0.40
V. xyele
fore leg
1.20
1.10
0.40
1.20
1.00
0.40
mid leg
3.40
1.90
1.10
3.50
1.90
1.10
hind leg
3.90
1.70
0.50
3.90
1.80
0.50
V. kurtokalami
fore leg
1.20
1.05
0.40
1.40
1.20
0.40
mid leg
3.30
1.90
1.20
3.70
2.15
1.35
hind leg
3.60
1.65
0.56
3.70
1.88
0.52
V. nieseri sp. n.
fore leg
1.25
1.05
0.36
1.35
1.14
0.40
mid leg
3.40
1.89
1.18
3.59
2.02
1.26
hind leg
3.61
1.50
0.50
3.86
1.77
1.57
V. heissi sp. n.
fore leg
1.11
0.98
0.37
mid leg
3.20
1.73
0.95
hind leg
3.19
1.37
0.48
V. imadatei
fore leg
1.30
1.00
0.34
1.18
1.10
0.36
mid leg
3.50
1.94
1.18
3.48
1.90
1.28
hind leg
3.69
1.40
0.43
3.57
1.36
0.48
V. yangae sp. n.
fore leg
1.15
1.02
0.39
1.00
0.96
0.36
mid leg
3.20
1.75
1.12
2.85
1.58
1.25
hind leg
3.28
1.30
0.48
3.30
1.44
0.44
Esakia ventidioides
fore leg
1.10
0.93
0.27
1.15
1.00
0.30
mid leg
3.20
2.44
0.92
3.60
2.50
0.95
hind leg
3.70
0.95
0.30
3.90
1.05
0.38
Table 3. Measurements of pronotum of macropterous form of
Ventidius species. For an explanation of the characters between
parentheses, see fig. 2.
Species Median humeral lateral lateral
length (j) width (k) length (1) length (m)
V. aquarius
2.70
2.00
0.80
1.90
V. usingeri
1.95
2.00
0.98
1.58
V. malayensis
2.10
2.25
0.10
1.60
V. henryi
1.32
1.42
0.60
1.18
V. hungerfordi
1.40
1.51
0.64
1.51
V. polhemorum
1.40
1.54
0.70
1.10
V. werneri
1.60
1.70
0.50
1.30
V. pilosus
1.65
1.80
0.77
1.38
V. modulatus
1.61
1.65
0.71
1.40
V. karen
1.48
1.90
0.30
0.84
V. pulai
1.45
1.66
0.40
1.25
V. xyele
2.55
1.70
0.75
1.35
V. xiphibion
2.70
1.75
0.88
1.45
V. kurtokalami
1.67
1.73
0.79
1.38
V. nieseri
1.75
1.90
0.80
1.40
208
J. H. EPLER', A. D. HARRISON2 & L. HARE3
' Crawfordville, U.S. A, 'Fish Hoek, South Africa, 'INRS-Eau, Quebec, Canada
ACINORETRACUS, A NEW AFROTROPICAL GENUS
FOR SOME SPECIES PREVIOUSLY PLACED IN
DICROTENDIPES (DIPTERA: CHIRONOMIDAE:
CHIRONOMINAE)
Epier, J.H., A.D. Harrison & L. Hare. 1999. Acinoretracus, a new Afrotropical genus for some
species previously placed in Dicrotendipes (Diptera: Chironomidae: Chironominae). - Tijd-
schrift voor Entomologie 141 [1998]: 209-220, figs. 1-33, tables 1-4. [ISSN 0040-7496]. Pub-
lished 1 March 1999.
A new genus, Acinoretracus, is established for four Afrotropical species previously placed in
Dicrotendipes. These species are Chironomus (Carteria) regalis Goetghebuer, 1936 (selected as
type-species for Acinoretracus); Ch. {Dicrotendipes) multispinosus Freeman, 1957; Ch. (D.)
penicillatus Freeman, 1957; and Ch. (D.) crispi Freeman, 1957. The adult male and female,
pupa and larva are described and figured for A. multispinosus and A. penicillatus, and notes are
given for the other two species. Acinoretracus 'is very close to Kiefferulusbm can be distinguished
by the following characters: adult male: superior volsella with dense apical brush of long, fine
setae, without large setae; inferior volsella with narrowed, upturned apex bearing several apical
setae; adult female: gonocoxite IX vestigial, without setae; pupa: cephalic tubercles low, round-
ed; thoracic horn base with anteriorly directed flange-like lobe; long, taeniate ventral an-
tepronotal and anterior precorneal setae; pedes spurii B and larval ventral tubules absent; larva:
S I plumose on inner side only; mentum with first and second lateral teeth not fused; short and
wide, medially contiguous ventromental plates, with basally forked striae; mandible with rugose
lateral margin and U shaped pecten mandibularis; lateral and ventral tubules absent.
Correspondence: Dr. J. H. Epler, 461 Tiger Hammock Rd., Crawfordville, FL 32327, USA. E-
mail: johneplr@freenet.tlh.fl.us
Key words. - Diptera; Chironomidae; taxonomy; Acinoretracus; Kiefferulus; Dicrotendipes;
Afrotropical.
Compared to the western Palaearctic and most of
the Nearctic, the taxonomy of the midge family Chi-
ronomidae from the Afrotropical region is poorly
known. Generic relationships of many species are un-
certain because their immature stages, often necessary
for delimiting genera, are unknown, in spite of their
ecological importance in African fresh waters. Here
we report on reared material of several problematic
species that require establishment of a new genus.
Goetghebuer (1936) placed a newly described
Afrotropical species, Chironomus regalis, in the sub-
genus Carteria Kieffer, 1921. Carteria had been shown
earlier (Strand 1928) to be preoccupied by Carteria
Diesing, 1866; Strand (1928) offered Carteronica as a
replacement name; Tendipes longilobus (Kieffer, 1916)
was retained as the type-species. Freeman (1957) con-
sidered Carteronica to be a synonym of Chironomus
{Dicrotendipes). Later workers, such as Hamilton et al.
(1969), re-elevated Dicrotendipes Kieffer, 1913, to
generic status. Epler (1987:19, 1988:9, 32) noted that
four Afrotropical species placed in Dicrotendipes by
Freeman & Cranston (1980) could not be maintained
in that genus. Epler (1988) also stated that Carteronica
was not a synonym of Dicrotendipes, and that Carteron-
ica longilobus did not belong with those Afrotropical
species previously considered to belong to Carteronica.
Cranston et al. (1990) showed that C longilobus was
best placed in Kiefferulus Goetghebuer, 1922; Car-
teronica became a junior synonym of Kiefferulus.
Based on the morphology of the adult genitalia and
the immature stages, the Afrotropical species 'Dicro-
tendipes multispinosus (Freeman, 1957) and 'D.' peni-
cillatus (Freeman, 1957) require establishment of a
new genus. Similarities in the adult genitalia, and to a
209
Tijdschrift voor Entomologie, volume i4i, 1998
lesser extent coloration, between these two species
and 'D'. crispi (Freeman, 1957) and 'D. ' regalis (Goet-
ghebuer, 1 936) allow the latter two species to also be
included in this grouping. In this paper, the new
genus name Acinoretracus is proposed for those four
Afrotropical species removed from Dicrotendipes. The
adult stages are redescribed, and the pupa and larva
described for the first time, for A. multispinosus and A.
penicillatus.
Methodology
Morphological terminology and abbreviations fol-
low Saîther (1980), Epier (1988) and Langton
(1994). For the anteromedian circular area of thinner
cuticle on the larval frontal apotome we adopt the
term apotomal fenestra, as suggested by Epler in
Cranston (1996). This structure is distinct from the
frontal pit found in Dicrotendipes, although some Di-
crotendipes (and other genera, such as Glyptotendipes)
may possess an apotomal fenestra (see Cranston 1 996
and Epler 1987, 1988).
Other abbreviations used: bmnh = British Muse-
um (Natural History).
Measurements are in urn, unless otherwise stated,
and consist of the range followed by the mean if more
than three specimens were measured. In the descrip-
tion of Acinoretracus multispinosus, data from Amakye
& Sasther (1993) are included in brackets ([]) if they
were outside of the range of the measurements per-
formed in this study; some data from their redescrip-
tion are also incorporated into the description of the
genus below.
Systematics
Acinoretracus Epler, Harrison et Hare gen. n.
Type-species: Chironomus (Carteria) regalis Goetghebuer,
1936: 465, by present designation.
Etymology
An anagram of Carteronica. Gender masculine.
Adult male
Medium-sized species, wing length about 1.6-3.0
mm; general colouration yellowish-brown to brown,
thorax with dorsal median dark stripe extending from
front to postnotum; abdomen with darker posterior
bands and/or median lines/triangles or almost com-
pletely dark; wings unmarked.
Head. — Eyes bare, with dorsomesal extension. Tem-
poral setae uniserial, beginning mesad to dorsomesal
eye extension and running behind the eyes. Antennae
with 1 1 flagellomeres; AR about 2.0-3.0. Frontal tu-
bercles minute/vestigial. Clypeus subquadrate, setose.
Cibarium with internal sensillae. Maxillary palp with 5
palpomeres, palpomere 1 weakly sclerotized; palpo-
mere 3 with 5-8 subapical sensilla clavata.
Thorax. - Antepronotum bare, lobes dorsally di-
vided. Scutum not extending over antepronotum,
scutal tubercle not present. Humeral pit obsolete;
thoracic scar moderately developed. Acrostichal setae
long, beginning close to antepronotum, 6-20; dor-
socentral setae 7-14/side, uniserial; prealar setae 4-
6/side; with one supraalar seta/side. Scutellum with
6-17 setae, uni- or biserial.
Wing. - Membrane without macrotrichia, with
moderate punctation of microtrichia. Brachiolum
with 2-3 setae and proximal and distal groups of sen-
silla campaniformia. Anal lobe well developed, apex
of wing rounded or slightly truncate. Veins R, Rl and
R4+5 with setae; squama with setae. Costa not ex-
tended. Apical and posterior subapical margin of
wing with scale-like setae.
Legs. - Apex of foretibia with rounded scale, with-
out spur; foretarsus without beard. Middle and hind
tibiae each with two combs; middle combs each with
1 spur; hind combs with inner comb with 1 spur, out-
er with 1-6 spurs. Sensilla chaetica present on apical
% of metatarsus of middle leg, sometimes with sensil-
la chaetica on metatarsus of hind leg. Tarsal claws
simple; empodium well developed; pulvilli small,
simple, about Vi length of claw.
Abdomen. - With moderate coverage of long setae,
arranged in loosely transverse rows.
Hypopygium (figs. 1, 8). - Anal tergal bands
strong, converging before anal point and continuing
onto point as a ridge. Anal point broad or narrow,
downturned apically and sometimes hooked. Median
anal tergite setae present or absent between dorsal
ridges at base of anal point, lateral setae present along
base of anal point. Superior volsella with short to elon-
gate cylindrical base, with dense apical brush of long,
fine setae, without large sensilla chaetica (setae); apex
sometimes bifid. Median volsella absent or present as
small wart-like protuberance that bears 4-10 long se-
tae. Inferior volsella with narrowed, upturned apex
bearing several apical setae, volsella sometimes swollen
dorsally before apex. Gonostylus semi-quadrate and
bulbous, sometimes with weak crista ventralis and a
moderately to well developed thinner outer heel, or
gonostylus more elongate and strongly curved medial-
ly, without crista ventralis or apical heel.
Adult female
As in the male, with following differences:
Head. - Antenna with 5 flagellomeres; AR about
0.33-0.46.
Thorax. - With about 2 humeral setae (anterior-
most dorsocentral setae).
Wing. - Slightly stouter than in male, with more
210
Epler et al.: Acinoretracus new genus
setae on veins.
Genitalia (figs. 11, 12). - Gonocoxite IX vestigial,
without setae. Gonapophysis VIII with well developed
dorsomesal and ventrolateral lobes. Apodeme lobe
well developed, with dense microtrichia. Labia with-
out microtrichia. Seminal capsules ovoid with a short
neck; spermathecal ducts without loops or bends.
Pupa
Exuviae brown, margins darker.
Céphalothorax. - Cephalic tubercles low, rounded;
frontal setae small (fig. 15). Dorsum mostly smooth,
with longitudinal row of tubercles, some of which are
sharply pointed. Thoracic horn plumose; base reni-
form, with anteriorly directed flange-like lobe, with 2
tracheal bundle scars (fig. 17). Dorsal antepronotal
seta short, ventral seta long and taeniate. Anterior
precorneal seta very long and taeniate, posterior seta
short (fig. 17). Four dorsocentral setae; Del and Dc2
close, Dc3 and Dc4 close; Del and Dc4 thicker than
Dc2 and Dc3.
Abdomen (fig. 18). -An uninterrupted row of pos-
terior hooklets on T II, about Vi width of segment. S
II and III with posterior transverse medial band of
long spines. Pedes spurii A present on S IV; pedes
spurii B absent. Segment VIII with dark caudolateral
combs, with 1-4 large spurs and 1-5 smaller spurs or
spines; without ventral tubules. Setation: Each side of
segment I with 1 lateral seta; II-IV with 3 lateral setae;
V-VII with 4 lateral taeniae, these arranged with first
two closer together on anterolateral margin and last
two closer together on posterolateral margin; VIII
with 5 lateral taeniae; anal lobe with a pair of dorsal
taeniae and a biserial fringe of about 80-130 taeni-
ae/side. Tergites and sternites with one pair of 0-se-
tae. Shagreen: T I bare; T II-V (VI) with a mostly
continuous field of points which become progressive-
ly larger posteriorly, weaker towards midline so that
posterior points appear in two groups, on T III-VI
these posterior shagreen groups slightly elevated
above rest of integument; T VI sometimes with ante-
rior and posterior fields of points, largest points in
posterior portion of posterior field; T VII-VIII with
anterior pair or transverse band of points, weaker on
T VIII; anal disc without shagreen. Conjunctives III-
IV, IV- V, V-VI with fine spinules. Pleura of II-III
(IV) with longitudinal bands of fine to coarse spin-
ules. S I bare; S II-V with scattered fine spinules; S
VI-VII (VIII) with anterior patches of fine spinules.
Larva
Head capsule yellow/yellow-brown, with darker
posterior margin, mentum, premandibles and mandi-
bular teeth. Two pairs of eyespots. Body usually
whitish in life but some specimens with red pigment;
claws brown.
Head. - Antenna (fig. 30) with 5 segments. Ring
organ in basal third of basal segment. Antennal blade
shorter than flagellum; accessory blade short, about Vò
length of segment 2. Style and Lauterborn organs pre-
sent at apex of segment 2.
Dorsum of head either with frontoclypeal apotome
and labral sclerite 2 (fig. 26), or frontal apotome and
labral sclerites 1 and 2, with anterior margin of
frontal apotome indistinct (fig. 23); apotome with
anteromedian fenestra. Labrum (fig. 28) with S I
plumose on inner side only; S II simple and on short
pedestal; S III simple; S IVA minute, 2-segmented; S
IVB simple, shorter than S IVA. Labral lamella with
marginal fringe. Pecten epipharyngis simple, with 9-
16 pointed teeth, no teeth on surface. Premandible
(fig. 31) with 5-6 teeth, brush well developed.
Mandible (fig. 32) with rugose outer margin, a pale
dorsal preapical tooth and dark apical and three inner
teeth. Pecten mandibularis composed of about 12
coarse setae arranged in U shape. Seta subdentalis
(fig. 33) apically widened with numerous fine apical
teeth. Seta interna plumose, with four main branches.
Mentum (figs. 24, 27) with 13 teeth, median tooth
trifid and lower than first lateral teeth; first lateral
teeth separate from seconds. Ventromental plates
short and wide, 3.5- 4. OX wider than long, contigu-
ous or nearly so medially; with smooth anterior mar-
gin; most striae complex, fork-like, with several
branches arising from base of each stria (fig. 25); in-
nermost and outermost striae simple. Setae submenti
simple, at base of ventromental plates. Maxilla (fig.
29) without serrate lacinial chaetae; maxillary palp
about twice as long as wide and with very long a seta.
Triangulum occipitale wide.
Body. — Anterior parapods with simple and pecti-
nate claws; posterior parapod claws simple. Lateral
and ventral tubules absent. Procerci slightly longer
than wide, each with 2 minute basal setae and 7-8
moderately long apical setae. Supraanal setae fine,
slightly shorter than anal tubules. Two pairs of anal
tubules, about 3X as long as wide.
Remarks
When all life stages are considered, Acinoretracus
can be seen to be very close to Kieffèrulus, not Axarus
or Lipiniella as suggested by Hare in Cranston, et al.
(1990). Kieffèrulus has recently been expanded by the
inclusion of several species previously assigned to oth-
er genera (Cranston et al. 1990); they (ibid.: 421)
presented an emended diagnosis for the genus. Note
that in their listing of included species, the authors
omitted the South African species K. nigropunctatum
(Freeman, 1957) (O. A. Ssther personal communica-
tion), K. modocensis (Sublette, 1960), a western Ne-
arctic species; and Epler (1995) recently moved Chi-
ronomus pungens (Townes, 1945), an eastern Nearctic
211
Tijdschrift voor Entomologie, volume mi, 1998
Figs. 1-7 Acinoretracus multispinosus, adult male. - 1, Hypopygium; 2, left superior volsella, dorsal; 3, left superior volsella,
lateral aspect of fig. 2; 4, left superior volsella, dorsal; 5, left superior volsella, dorsal; 6, left superior volsella of holotype, dor-
sal; 7, inferior volsella, lateral.
species, to Kiejferulus. The diagnosis of Cranston et
al. (1990) is already in need of further emendation,
for, as noted below, an undescribed southern Nearc-
tic species of Kiefferulus has been found in which the
larval ventromental plates are contiguous medially.
One of us (ADH) is not in complete agreement with
their synonymy of some African Nilodorum species
with Kiejferulus.
While some life stages (the female) of Acinoretracus
may not be clearly separable from Kiefferulus as cur-
rently defined, the complete suite of characters taken
from all life stages demonstrates the generic unique-
ness of Acinoretracus from Kiefferulus. These differ-
ences include:
Adult male: superior volsella with dense apical
brush of long, fine setae, without large setae; inferior
volsella with narrowed, upturned apex bearing sever-
al apical setae. No described species of Kiefferulus pos-
sesses such genitalia. Two species, A. multispinosus
and A. regalis, possess a rudimentary median volsella
that bears several large setae.
Adult female: gonocoxite IX vestigial, without se-
tae. Sasther (1977: 170) describes the female genitalia
of Kiefferulus with gonocoxite IX 'small, with about 2
setae'. However, Cranston et al. (1990: 423) noted
that in K longilobus gonocoxite IX is small and ap-
parently without setae; while Harrison (1996: 10)
found 'gonocoxite IX large with about 1 0 setae' in K.
cbloronotus (Kieffer).
Pupa: cephalic tubercles low, rounded; thoracic
horn base with anteriorly directed flange-like lobe;
long, taeniate ventral antepronotal and anterior pre-
corneal setae; pedes spurii B and larval ventral tubules
absent. Note that while in Holarctic Kiefferulus the
precorneal setae are subequal, in the Afrotropical
212
Epler et al.: Acinoretracus new genus
species K. fractilobus (Kieffer), the posterior pre-
corneal seta is much larger and taeniate (JHE, unpub-
lished data based on rearings from Nigeria by LH).
Larva: S I plumose on inner side only; mentum
with first and second lateral teeth not fused; short and
wide, contiguous ventromental plates, with forked
striae; mandible with rugose outer margin and U
shaped pecten mandibularis; ventral tubules absent.
Pinder & Reiss (1983) and Cranston et al. (1990) di-
agnose Kiefferulus larvae as having medially separated
ventromental plates. However, an undescribed south-
ern Nearctic species {Kiefferulus sp. A in Epler 1992,
1995) has contiguous ventromental plates. This
species has been reared by JHE and has a pupa and
adult very similar to K dux (Johannsen). Note that
the ventromental plate striae of K sp. A are simple,
not forked as in Acinoretracus (fig. 25).
It can be hypothesized that Acinoretracus and Kief-
ferulus form a sister group within the Chironomus
group (as defined by Epler 1988: 194), but the nature
of the relationships between Kiefferulus and related
genera must await a world-wide revision of Kiefferu-
lus, utilizing characters from all life stages.
Acinoretracus multispinosus (Freeman) comb. n.
(figs. 1-7, 14, 17-22, 23-25)
Chironomus {Dicrotendipes) multispinosus Freeman 1957:
373 (original description of adult male).
Dicrotendipes multispinosus (Freeman). Freeman &
Cranston 1980: 190 (catalog).
'Carteronica' multispinosus Freeman. Cranston et al. 1990
(larval ventromental plates).
Dicrotendipes multispinosus (Freeman). Amakye & Ssether
1993 (redescription of adult male; description of adult fe-
male).
Description
The male of this species was recently redescribed by
Amakye & Ssether (1993); they also described the fe-
male in detail. Some of their data are included below
in brackets in the descriptions of those life stages.
Adult male (n=4-5). - Colour. Head yellowish-
brown, antennae light brown; thorax yellowish with
dark median stripe extending from front of scutum to
postnotum; wings unmarked, pale yellowish-brown;
haltere pale; legs yellowish with femoral apices brown,
bases and apices of tibiae brown; abdomen yellowish
with brown posterior triangular areas/bands, tergite
VIII and hypopygium almost completely brown.
Length. Total 3.63-4.60, 4.21 mm [3.31-4.48,
3.98 mm]; thorax 0.90-1.03, 1.02 mm; abdomen
2.73-3.58, 3.19 mm.
Head. Frontal tubercles 2.5 long. Temporal setae
12-19; clypeal setae 12-23, 18; cibarial sensilla 4-8.
Lengths of palpomeres 1-5: 43-50 [30-56, 44]; 45-47
[37-52,47]; 128-150 [135-168, 152]; 145-165 [150-
179, 165]; 200 [233-248, 235]. AR 2.79-2.78 [2.28-
2.60, 2.48].
Thorax. Setae: acrostichals 7- 18 [6-9, 8]; dorsocen-
trals 10-11, 10 [7-14, 9]; prealars 5 [4-5, 5]; scutellars
8-16 [6-12, 9].
Wing. Wing length 1.63-2.10, 1.86 mm [1.74-
2.02, 1.88 mm]; width 0.44-0.61 mm. VR 1.16-
1.24. Setae: R 20-34; Rl 17-26; R4+5 20-39; squama
4-9 [6-10]. Wing apex rounded (fig. 14).
Legs. Tarsomere 1 of middle leg with 7-10 sensilla
chaetica; tarsomere 1 of hind leg with 0-1 sensilla
chaetica. Lengths and proportions of legs see Table 1.
Hypopygium (fig. 1). Superior volsella cylindrical,
shaft bare but with expanded pad with dense brush of
long setae apically (figs. 4, 5), or apex bifid (figs. 2, 3,
6). Median volsella reduced to small protuberance,
with [4] 5- 10, 7 large anteromedially directed setae.
Inferior volsella with narrowed, upturned apex, bear-
ing 3-7 large setae and several smaller setae; volsella
dorsally expanded preapically (fig. 7). Gonostylus
bulbous/quadrate, with crista ventralis. Anal point
ridge bearing 2-5 setae; anal point with 8-12 smaller
lateral setae.
Adult female (n=l). - Colour. Similar to male.
Length. Total 4.00 mm [4.62 mm]; thorax 1.15
mm; abdomen 2.85 mm.
Head. Frontal tubercles 2.0 long. Temporal setae
16; clypeal setae 18; cibarial sensilla 14. Lengths of
palpomeres 1-5: 45 [56]; 52 [56]; 150 [180]; 155
[184]; 250 [263]. AR 0.46 [0.45].
Thorax. Setae: acrostichals 14; dorsocentrals 11+2
humerais [16], 10; prealars 5; scutellars 15 [13].
Wing. Wing length 1.90 mm; width 0.65 mm. VR
1.17 [1.19]. Setae: R31;R1 31; R4+5 42; squama 14 [12].
Legs. Tarsomere 1 of middle leg with 19 sensilla
chaetica; tarsomere 1 of hind leg with 10 sensilla
Table 1 . A. multispinosus male: lengths and proportions of
legs (n = 1- 2).
PI
P2
P3
fe
[942] 1060-1160
[887] 1000-1070
[867] 1020-1100
ti
[638] 740-810
[742] 890
[943] 1100-1150
tal
[1420] 1480
[457] 520
750
ta2
[667] 690
[220] 230
400-420
ta3
[581] 610
[157] 170-180
320-330
ta4
570
[95] 110-120
175-190
ta5
240
[57-67] 85-100
100-120
LR
2.00 [2.22]
0.58-0.63 [0.71]
0.68-0.73
BV
1.55
3.83-4.24
2.76-3.04
SV
1.22 [1.15]
[3.35] 3.50-3.63
2.68-2.83
213
Tijdschrift voor Entomologie, volume mi, 1998
Figs. 8-13. Acinoretracus penicillatus, adult male and female. - 8, Hypopygium; 9, superior volsella; 10, inferior volsella, lat-
eral; 11, female genitalia, ventral; 12, female genitalia, lateral; 13, wing apex. - Fig. 14. A. multispinosus, wing apex.
214
Epler et al.: Acinoretracus new genus
chaetica. Lengths and proportions of legs see Table 2.
Abdomen. Notum 150 [128] long. Tergite X with
8 setae. Cercus 125 [145] long.
Pupa (n=4-6). - Colour. Exuviae light brown, with
clear band at posterior margin of T (II) III-V, T VI-
VIII and anal lobes paler brown.
Length. Total 4.55-5.52, 5.10 mm; céphalothorax
1.25-1.38, 1.33 mm; abdomen 3.30-4.15, 3.78 mm.
Céphalothorax. Frontal setae 30-35, 32 long. Dor-
sal antepronotal seta 80-100 long, thin (n=2); ventral
antepronotal seta at least 113 long, taeniate (n=l).
Anterior precorneal seta around 195-238 long, taeni-
ate; posterior seta about 15 (broken?). Dorsocentral
setae lengths (n=l): Del 48; Dc2 83; Dc3 45; Dc4
33. Dorsum with row of 7-1 1, 9 tubercles.
Abdomen (fig. 18). T II with posterior row of 22-
27, 25 hooklets. S II (fig. 19) with posteromedian row
of 33-54, AG spines; S III with posteromedian row of
3-15, 10 spines. Shagreen on T II- VI with spinules
larger anteromedially, followed posteriorly by smaller
spinules which become progressively larger posterior-
ly, with posterior groups of larger spines separated
medially by area of much smaller spinules; shagreen
area on T II consists of a small rectangular area and
two anterolateral patches, these patches sometimes
joined with median patch to form broad T-shaped
area; on T III-V shagreen broadly quadrilateral or X-
shaped, with small fenestrations; on T VI broadly tri-
angular; on T VII with two anterolateral patches or
patches joined to form band; T VIII with two small
anterolateral patches of fine spinules or sometimes
with two longitudinal bands of fine spinules. T VIII
with 1-3 large and several smaller caudolateral spurs
(figs. 20-22). Anal lobes with 96-128, 113 taeniae.
Larva (n=4-5). - Colour. Head capsule light yel-
low-brown.
Head. Postmentum length 185-202, 191. Frontal
apotome and labral sclerite 1 not fused but anterior
margin of apotome not distinct, apotome weakly peb-
bled (fig. 23). Length of antennal segments 1-5: 68-
85, 76; 22-30, 28; 11-15, 14; 7-8, 8; 5-7, 6; AR 1.08-
1.40, 1.26. Premandible 80-93, 86 long. Pecten
epipharyngis with 13 teeth (n=2). Mandible length
158-172, 167; pecten mandibularis with 11-14, 13
setae. Mentum (fig. 24) width 1 15-125, 121. Ventro-
mental plates 158-175, 166 wide; 43-47, 45 long;
VPR 3.51-3.84, 3.73; with 41-52, 48 striae.
Remarks
Freeman (1957: 373) stated the superior vulsella of
multispinosus was bifid. However, in many specimens
the apex is not bifid, but pad-like with a dense brush
of setae (figs. 4, 5; see also Amakye & Sa»ther 1993:
figs. 2F, 2J, 2K). In the holotype specimen and three
Table 2. A. multispinosus female: length and proportions of
legs (n=l).
PI
P2
P3
fe
[1040] 1060
[973] 1000
[983] 1040
ti
720 [756]
[860] 880
[1068] 1080
tal
1460
480
690
tal
630
210
370
ta3
560
160
310
ta4
560
110
190
ta5
220
100
115
LR
2.03
0.55
0.64
BV
1.64
4.07
2.85
SV
1.22
3.92
3.07
of the five males associated with immature stages from
Nigeria, one a pharate male still within its pupal skin,
the apex of the volsella is bifid (figs. 2, 3, 6). In anoth-
er pharate male pupa, the superior volsellae were
broadly setose, similar to fig. 4. No other differences
were noted among the adults, and no differences were
noted in the immature stages between those adults
with apically bifid or pad-like superior volsellae.
In Acinoretracus multispinosus and A. regalis a small,
median protuberance, which bears several large setae,
is present above the base of each inferior volsella.
Amakye & Saldier (1993) termed the protuberance a
median volsella; this structure is absent in A. crispi
and A. penicillatus.
Amakye & Saldier (1993: 269, 272) refer to a 'more
or less distinct crista dorsalis' on the gonostylus of mul-
tispinosus. However, this ridge-like structure is actually
ventral and could be termed a crista ventralis. This
structure is also present on A. regalis and A. penicillatus.
See Remarks under A. regalis for comments concern-
ing the separation of A multispinosus and A. regalis.
Pupae of this species and A. penicillatus may prove
difficult to separate. In general, the pupa of A. peni-
cillatus'^ larger (5.53-5.91 mm vs. 4.55-5.53 mm in
A. multispinosus) and has a higher T II hooklet count
(26-30 vs. 22-27 in A. multispinosus) but fewer anal
lobe taeniae (84-100 vs. 96-128 in A. multispinosus).
These numbers may prove meaningless when more
populations are sampled; our sample is admittedly
small. The shagreen spinules on T II-VI are larger an-
teromedially in A. multispinosus; in A. penicillatus
they are subequal anteromedially. Another useful
character for separation is the presence of larger ante-
rior and posterior triangular clear areas on T VII-VIII
on A. penicillatus; such areas are reduced or not pre-
sent on our A. multispinosus material.
Larvae of A. multispinosus are very similar to A.
penicillatus. In our material, fourth instar A. multi-
spinosus larvae were smaller; however size differences
may not hold throughout the species' range and in
general may not provide a good character for separa-
215
Tijdschrift voor Entomologie, volume i4i, 1998
19
If f] ]
*m
Figs. 15-16. Acinoretracus penicillatus pupa. - 15, Cephalic tubercles; 16, tergite Vili caudolateral spurs. - Figs. 17-22. A.
multispinosus, pupa. - 17, Thoracic horn base and precorneal setae; 18, abdomen, dorsal; 19, sternite II; 20-22, tergite VIII
caudolateral spurs, variations.
216
Epler et al.: Acinoretracus new genus
tion. A good distinguishing feature is the well defined
labral sclerite 1 of A. multispinosus; in A. penicillatus
this sclerite is fused with the frontal apotome.
The immature stages of A. multispinosus described
in this paper were collected from Lake Opi, a shallow
(maximum depth 4 m), slightly acidic (pH 6.0-6.5)
and dilute (low in dissolved minerals) body of water in
the West African Guinea Savanna region near Nsuk-
ka, Nigeria. Larvae of A. multispinosus were uncom-
mon at depths greater than 0.5 m in this lake (Hare &
Carter 1986). For more detailed information on the
chemistry of Lake Opi see Hare & Carter (1984).
The species is known from Burkina Faso, Came-
roon, Chad, Ghana, Nigeria, Uganda and Zaire.
Material examined. - [burkina faso] Haute Volta,
a.o.f., Tangrela, 3-12-56, Cercle de Banfora, J. Ha-
mon, ORSTOM ree, 1 6 paratype (bmnh). Cameroon:
Kumba, vi-vii-58, D. J. Lewis, 1 â (bmnh). Nigeria:
Lake Opi nr. Nsukka, 8-iii-1977 to 6-viii-1980, leg. L.
Hare, numerous associated adults, pupal exuviae and
larvae, uganda: Lake Bungani, l-viii-1932, G. H. E.
Hopkins, holotype S (bmnh).
Acinoretracus penicillatus (Freeman) comb. n.
(figs. 8-13, 15, 16,26-33)
nee Carteria regalis Goetghebuer. Freeman 1955: 371. [mis-
identification].
Chironomus (Dicrotendipes) penicillatus Freeman 1957: 374
(original description of adult male).
Dicrotendipes penicillatus (Freeman). Freeman & Cranston
1980: 190 (catalog).
Description
Adult male (n=3). - Colour. Head and antennae
light brown; thorax brown, with dark median stripe
extending from front of scutum to postnotum; legs
brown; wings unmarked, pale brownish; abdomen
brown, tergites I-VI with central dark stripe, VII,
VIII and hypopygium uniform brown.
Length. Total 4.15-4.90 mm; thorax 1.05-1.15
mm; abdomen 3.10-4.90 mm.
Head. Frontal tubercles 2.0 long. Temporal setae
11-15; clypeal setae 18-22; cibarial sensilla 11-18.
Lengths of palpomeres 1-5: 30-33; 42-45; 108-113;
105-125; 154-170. AR 1.95-2.08.
Thorax. Setae: acrostichals 11-13; dorsocentrals
10-11; prealars 5-6; 7-8.
Wing. Wing length 2.00-2.03 mm; width 0.50-
0.54 mm. VR 1.16-1.20. Setae: R 25-30; Rl 13-19;
R4+5 20-23; squama 6-7. Wing apex slightly trun-
cate (fig. 13).
Legs. Tarsomere 1 of middle leg with 8-13 sensilla
chaetica; tarsomere 1 of hind leg with 0 sensilla chaet-
ica. Lengths and proportions of legs see Table 3.
Hypopygium (fig. 8). Superior vulsella digitiform,
shaft bare but with dense brush of long setae apically
(fig. 9). Median volsella absent. Inferior vulsella with
narrowed, upturned apex, bearing 3-4 large setae and
several smaller setae (fig. 10). Gonostylus bulbous /qua-
drate, with crista ventralis. Anal point ridge without
dorsal setae; anal point with 3-6 smaller lateral setae.
Adult female (n=l-2). - Colour. Similar to male.
Length. Total 3.77-4.26 mm; thorax 1.12-1.28
mm; abdomen 2.65-2.98 mm.
Head. Frontal tubercles vestigial. Temporal setae
13-17; clypeal setae 22-32; cibarial sensilla 17-18.
Lengths of palpomeres 1-5: 38-40; 40-43; 100-112;
108-117; 177-185. AR 0.33.
Thorax. Setae: acrostichals 13-14; dorsocentrals
10-11 +2 humerais; prealars 5; scutellars 8.
Wing. Wing length 2.18-2.35 mm; width 0.63-
0.70 mm. VR 1.14-1.16. Setae: R 31; Rl 20-23;
R4+5 35-41; squama 9-10.
Legs. Tarsomere 1 of middle leg with 21-23 sensil-
la chaetica; tarsomere 1 of hind leg with 0 sensilla
chaetica. Lengths and proportions of legs see Table 4.
Abdomen. Notum 155 long. TergiteX with 10-11
setae. Cercus 113 long.
Pupa (n=3-4). - Colour. Exuviae brown, with clear
band at posterior margin of T II-VI, T VII- VIII with
clear anterior and posterior triangular areas; anal
lobes paler brown.
Table 3. A. penicillatus male: lengths and proportions of legs
(n = 2-3):
PI P2 P3
fe
990-1040
920-950
1020-1040
ti
745-760
860-880
1140-1160
tal
1130-1150
400-420
690-700
ta2
530-540
220-240
410-420
ta3
475-480
190-195
330-340
ta4
380
120
190
ta5
180-185
95-100
115-120
LR
1.49-1.54
0.47-0.48
0.60-0.61
BV
1.83-1.86
3.38-3.60
2.71-2.73
SV
1.55
4.36-4.45
3.13-3.14
Table 4. A. penicillatus female: lengths and proportions of
legs (n=2-3).
PI P2 P3
fe
1000
960
1000-1090
ti
760
910
1120-1220
tal
1250
460
680-710
ta2
540
230
345-400
ta3
460
190
320-330
ta4
360
120
180
ta5
180
100
120
LR
1.64
0.51
0.58-0.61
BV
1.95
3.64
2.90-2.93
SV
1.41
4.07
3.12-3.25
217
Tijdschrift voor Entomologie, volume i4i, 1998
Figs. 23-25. Acinoretracus multispinosus larva. - 23, Apotome and labral sclerites; 24, mentum and ventromental plate; 25, de-
tail of ventromental plate striae. - Figs. 26-33. A. penicillatus, larva. - 26, Apotome and labral sclerites; 27, mentum and ven-
tromental plate; 28, labral structures; 29, maxilla; 30, antenna; 31, premandible; 32, mandible; 33, seta subdentalis.
218
Epleretal.: Acinoretracus new genus
Length. Total 5.53-5.95 mm; céphalothorax 1.30-
1.38 mm; abdomen 4.20- 4.65 mm.
Céphalothorax. Frontal setae 30-50, 38 long. Dor-
sal antepronotal seta not measurable (broken off?);
ventral antepronotal seta at least 1 50 long, taeniate (n
= 1). Anterior precorneal seta not measurable, taeni-
ate; posterior seta about 80. Dorsocentral setae
lengths (n=2-3): Del 40-50; Dc2 100; Dc3 21-33;
Dc4 40-43. Dorsum with row of 1 1 - 1 7, 13 tubercles.
Abdomen. T II with posterior row of 26-30, 28
hooklets. S II with posteromedian row of 27-36, 33
spines; S III with posteromedian row of 10-15, 13
spines. Shagreen on T II-VI becomes progressively larg-
er posteriorly, with posterior groups of larger spines
separated medially by area of smaller spinules; on T II
consists of broad T- or V-shaped area; on T III- T V
shagreen broadly quadrilateral or X-shaped, with small
fenestrations; on T VI broadly triangular; on T VII
with two anterolateral patches or patches joined to form
band; T VIII with two anterior patches of fine spinules.
T VIII with 3-4 large and several smaller caudolateral
spurs (fig. 16). Anal lobes with 84-100, 92 taeniae.
Larva (n=3). - Colour. Head capsule yellow-brown.
Head. Postmentum length 235-260. Frontal apo-
tome and labral sclerite 1 fused, apotome strongly
pebbled (fig. 26). Length of antennal segments 1-5:
75-90; 28; 13-15; 10; 5; AR 1.19-1.48. Premandible
85-100 long. Pecten epipharyngis with 14-16 teeth
(n=2). Mandible length 175-190; pecten mandibu-
laris with 8-12 setae. Mentum (fig. 27) width 123-
135. Ventromental plates 188-203 wide; 50-54 long;
VPR 3.57-3.76; with 48-50 striae.
Remarks
See remarks under A. multispinosus for separation
of the immature stages.
Our associated material was collected at the type-lo-
cality from bottom mud in acid, brown water dys-
trophic lakes and ponds at a depth of about 1.5 m; they
were not collected in the marginal reeds. The small
lake at Betty's Bay is described by Harrison (1958).
The species is known only from South Africa.
Material examined. - [south africa]: Cape
Province, Cape Peninsula, Hout Bay, Skoorsteenskop,
11-1951, P. Brinck, 1 cT paratype (bmnh). south
africa: Western Cape Province: Betty's Bay, Malgat
Vlei, January and February 1994, leg. A.D. Harrison,
numerous associated adults, pupal exuviae and larvae.
Acinoretracus regalis (Goetghebuer) comb. n.
Chironomus (Carteria) regalis Goetghebuer 1936: 465 (orig-
inal description of adult male).
Chironomus {Dicrotendipes) regalis Goetghebuer. Freeman
1957: 373 (redescription).
Dicrotendipes regalis (Goetghebuer). Freeman & Cranston
1980: 190 (catalog).
This species is very similar to A. multispinosus, dif-
fering mainly in having a normal spur count on the
hind tibial combs. Freeman (1957) stated that the two
species could be separated by differences in the geni-
talia, such as the anal point being narrower in side
view in multispinosus (this difference has been difficult
to observe), more rounded gonosrylus in multispinosus
(Freeman himself stated that this difference may not
be constant), and the bifid nature of the superior
vulsella in multispinosus (which, as discussed above, is
not always bifid). The two species are so similar in
structure that it may be that both are the same; A.
multispinosus may only be a variety with extra spurs on
the hind tibial comb. We have made regalis the type-
species for the genus because it was the first described
species; should multispinosus fall as a synonym, no
name changes or change in type-species status would
be necessary. Reared material and more adult material
will be needed to solve the potential problem of sepa-
ration of regalis from multispinosus. The immature
stages ofyl. regalisele unknown.
The species is recorded from Burkina Faso, Ghana,
Sierra Leone and Zaire.
Material examined. - [ghana] Gold Coast: Addah,
Adidome, 1921, N.L. Braybrock, in hut, 2 cT
(bmnh). [zaire]: Congo-Beige: Eala, 17-iv-1936, J.
Ghesquière (label reads 'compared with holotype by
P. Freeman 1955'), 1 cT (bmnh).
Acinoretracus crispi (Freeman) comb. n.
Chironomus {Dicrotendipes) crispi Freeman 1957: 374 (orig-
inal description of adult male).
Dicrotendipes crispi (Freeman). Freeman &C Cranston 1980:
190 (catalog).
This species differs from the other three in the
genus by having a more slender and strongly curved
gonostylus, long and slender superior volsella, and a
narrow anal point that is sharply hooked apically
(Freeman 1957: figs. 8g, 8j). The inferior volsella is
similar to that of A. penicillatus in that it lacks the
preapical dorsal swelling. The immature stages are
unknown.
The species is recorded from Chad, Ghana, Mali,
Nigeria and Sudan.
Material examined. — [ghana]: Gold Coast: Red
Volta, Nangodi, 8-X-1954, G. Crisp, 1 S paratype
(bmnh).
219
Tijdschrift voor Entomologie, volume hi, 1998
Key to adult males of Acinoretracus
1 . Gonostylus broad, bulbous, semi-quadrate; superi-
or volsella short ; anal point broad (figs. 1, 8) 2
- Gonostylus more slender, strongly curved; supe-
rior vulsella long and slender; anal point slender
(see Freeman 1957: figs 8f, 8j) A. crispi
2. Hind tibia with 1 spur on outer tibial comb ....3
- Hind tibia with 4-5 spurs on outer tibial comb...
A. multispinosus
3. Median vulsella present with 4-7 large setae; infe-
rior vulsella with dorsal subapical swelling (simi-
lar to figs. 1, 7; see also Freeman 1957: figs. 8d,
8h) A. regalis
- Median volsella absent; inferior volsella without
subapical swelling or at most a slight dorsal ex-
pansion (figs. 8, 10) A. penicillatus
Acknowledgments
We are grateful to Drs. P. S. Cranston and S. J.
Brooks for the loan of material from the British Mu-
seum (Natural History) and to Mr. B. A. Caldwell,
Dr. Cranston and Dr. O. A. Sasther for their com-
ments on the manuscript. The senior author wishes
to especially thank Dr. Barry Merrill and Judy Merrill
(Merrill Consultants, Dallas, TX) for providing funds
for laboratory and computer equipment.
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Received: 29 September 1998
Accepted: 12 November 1998
220
Hans R. FEIJEN
National Museum of Natural History, Leiden
A REVISION OF EURYDIOPSIS FREY (DIPTERA,
DIOPSIDAE) WITH DESCRIPTION OF FOUR NEW
ORIENTAL SPECIES
Feijen, Hans R., 1999. A revision of Eurydiopsis Frey (Diptera, Diopsidae) with description of
four new Oriental species. - Tijdschrift voor Entomologie 141 [1998]: 221-240, figs. 1-72,
table 1. [issn 0040-7496]. Published 1 March 1999.
Eurydiopsis Trey is defined. Catalogue and key to the six species are given, while differential and
quantitative characters are discussed. The earlier proposed synonymy between E. subnotata
(Westwood) from the Philippines and E. argentifera (Bigot) from Indonesia and Malaysia is re-
jected. Both species are redescribed, whereas E. helsdingeni sp. n. from Indonesia (Java), E.
glabrostylus sp. n. from the Philippines, E. brevispinus sp. n. from Laos and Myanmar and E.
sarawakensis sp. n. from Malaysia (Sarawak) are described.
H. R. Feijen, P. O. Box 639, Thimphu, Bhutan or (permanent) P. O. Box 2068, 2301 CB Lei-
den, The Netherlands
Keywords. - Diopsidae; Eurydiopsis; catalogue; key; new species; redescriptions; Asia.
In 1928 Frey erected the subgenus Eurydiopsis in
Diopsis Linnaeus 1775. As type-species Diopsis subno-
fczta Westwood 1848 from the Philippines was desig-
nated. The only differential character indicated by
Frey was constituted by the short and blunt facial
teeth of the Oriental Diopsis subnotata and 'Diopsis
circularis Macquart' (sensu van der Wulp 1897), as
opposed to the long and sharp facial teeth of the
African Diopsis. However, many African Diopsis do
not have facial teeth and it has been pointed out sev-
eral times (e.g., Shillito 1971: 289, Feijen 1989: 14)
that presence or absence of facial teeth cannot be used
as a key character to distinguish a genus.
Feijen (1978) showed that Diopsis circularis Mac-
quart 1835 represents an exclusively African species.
The species identified as Diopsis circularis by van der
Wulp (1897) and called Eurydiopsis circularis by Frey
(1928) had to represent an undescribed Oriental
species. Bigot (1874) described Diopsis argentifera
from Celebes (Sulawesi), Indonesia. However, in
1881 Bigot placed his Diopsis argentifera in synonymy
with Diopsis subnotata, quoting Osten Sacken who
published the synonymy in 1882.
Séguy (1949) described two Malagasy diopsids as
Diopsis {Eurydiopsis) apollo and Diopsis {Eurydiopsis)
apographica. Later, Vanschuytbroeck (1965) de-
scribed two more Malagasy species as Eurydiopsis an-
jahanaribei and Eurydiopsis vadoni. Shillito (1971)
thought it necessary to erect a new Malagasy genus
for these four 'Eurydiopsis, but Steyskal (1972) placed
all four Malagasy 'Eurydiopsis in Cyrtodiopsis. Cogan
& Shillito (1980) agreed to the inclusion in Cyrtodi-
opsis. Feijen (1981), in a note on Cyrtodiopsis, consid-
ered Shillito's original view correct, while Feijen
(1989) included the four species in Eurydiopsis S. &
V. nee Frey, a genus still to be described. The Mala-
gasy species can, for instance, be distinguished from
Eurydiopsis by their distinct inner vertical bristle (usu-
ally on a cone) and the non-articulated gonostyli with
an apodeme.
Burkhardt & de la Motte (1985: 205) expressed the
view that Cyrtodiopsis quinqueguttata was more relat-
ed to Eurydiopsis argentifera (as subnotata) than to any
Cyrtodiopsis. Feijen (1989) agreed that quinqueguttata
is an aberrant species, but doubted that it is more re-
lated to Eurydiopsis than to the other Cyrtodiopsis.
Steyskal (1972) elevated Eurydiopsis to genus level,
while Feijen (1989) stated that a redescribed Eury-
diopsis should be maintained as a separate genus. So
far, this genus only counted the one recognized Ori-
ental/Australasian species, but study of material from
various countries showed subnotata to represent a
species complex.
The material identified by van der Wulp (1897:
189) as ' circularis 'was rediscovered in the Leiden Mu-
seum and the Brussels Museum. It proved to represent
an undescribed Eurydiopsis, clearly distinct from the
subnotata complex. This new species is now described
221
Tijdschrift voor Entomologie, volume i4i, 1998
as E. helsdingeni. The synonymy of E. argentifera with
E. subnotata is rejected and argentifera is now recog-
nized as a distinct species occurring in Sulawesi, Java,
Sumatra and peninsular Malaysia. Three more species
were recognized in the subnotata complex: E. glabrosty-
lus sp. n. as the second species from the Philippines, E.
brevispinus sp. n. from Laos and Myanmar and E.
sarawakensis sp. n. from Sarawak, Malaysia.
A diagnosis of Eurydiopsis, a catalogue and a key to
the six species are presented. The geographic distribu-
tion of the Eurydiopsis species is briefly discussed. The
phylogeny and biology of the genus is reviewed. The
phylogenetic position of the genus Eurydiopsis in the
Diopsinae is not yet clear. A possible closer relation-
ship with the Diopsis indica complex is summarily in-
dicated. Various differential and quantitative charac-
ters are discussed.
Abbreviations used
IVB
Inner Vertical Bristle
1/w
OVB
length/width
Outer Vertical Bristle
SC
scutellar
scsp
SE
scutellar spines
Standard Error
tub
tubercles
Acronyms for collections
bmnh
BPBM
IRSNB
UZMH
ZMA
British Museum (Natural History), London
Bernice P. Bishop Museum, Hawaii
Institut Royal de Sciences Naturelles de
Belgique, Bruxelles
Wissenschaftsbereich Zoologie, Sektion
Biowissenschaften, Martin-Luther-
Universität, Halle
Nationaal Natuurhistorisch Museum,
Leiden (formerly Rijksmuseum van
Natuurlijke Historie)
Hope Entomological Collections, University
Museum, Oxford
Zoological Museum, University of Helsinki
Zoologisch Museum, Amsterdam
Genus Eurydiopsis Frey 1928
Eurydiopsis Frey 1928: 70 (as subgenus of Diopsis, type
species subnotata Westwood by original designation);
Shillito 1971: 288 (as subgenus); Steyskal 1972: 10 (as
genus); Steyskal 1975: 33 (as genus); Feijen 1989: 62 (as
genus).
Diagnosis
Large Diopsidae (7-13 mm) with relatively very
small eye stalks (eye span less than % of body length);
bipartite arista; medium-sized ovb, rvB a minuscule
wart or a small bristle; facial sulcus present; facial teeth
absent or very small and blunt (Feijen 1989 mentions
'no facial teeth' for Eurydiopsis in his key to the Diop-
sidae genera. This should be replaced by 'facial teeth
absent or very small and blunt'); precoxal bridge pre-
sent; supra-alar and infra-alar spines absent; scutal
bristles absent; scutellar spines medium-sized to long,
straight to simply curved; apical bristle absent; meta-
pleural spines strong and laterally directed; thorax uni-
formly pollinose; front femora slender to incrassate
with two long rows of tubercles, the outer row count-
ing on the average 19-23 tubercles and the inner row
22-26; no apical spurs on femora 2 and 3; mid tibiae
with two small subapical bristles; alula absent, sixth
vein not extending beyond anal cell, fifth vein extend-
ing beyond posterior crossvein but not reaching wing
margin; wings patterned with two rows of hyaline
spots and hyaline apex or preapical band; abdomen
moderately to strongly davate; syntergum consisting
of terga 1-3 or terga 1-4, seam between terga 2 and 3
distinct; spiracle 1 in tergum; female tergum 8 divided
or not, female tergum 10 with 3 to 4 pairs of hairs; fe-
male spiracle 7 in membrane, female sternum 8 a sin-
gle plate, sometimes strongly constricted medially;
three smooth, rounded spermathecae; male sternum 5
divided or single; male sternum 6 divided; male ster-
num 7+8 a very short, band-like sclerite with spiracle
7 located anteriorly of it (fig. 26); periandrium short,
rounded; gonostyli simple, fused to periandrium but
appearing articulated; processus longi fused (running
from gonostylus to gonostylus via edge of periandri-
um); phallapodeme simple; ejaculatory apodeme
small; no sexual dimorphism in length of eye stalks
and shape of front femora.
Catalogue
Eurydiopsis Frey, 1928: 70 (as subgenus of Diopsis).
Type species subnotata Westwood 1848, by
original designation).
Ref.: Steyskal 1972: 10; Steyskal 1975: 33;
Feijen 1989: 62.
argentifera Bigot, 1874: 112 {Diopsis).
Type locality: Celebes (Sulawesi), Indonesia.
Ref.: Bigot 1881: 373 (synonymy with
'subnitidd Westwood, quoting Osten Sacken
1882). Two syntypes (as 'cotypes') in umo.
brevispinus sp. n.
Type locality: Mt. Victoria, Myanmar. Holotype
and 1 paratype in bmnh, 3 paratypes in mluh.
glabrostylus sp. n. Type locality: Aroroy, Philippines.
Holotype and 9 paratypes in uzmh. Ref.: Frey
1928: 71 (in part, as subnotata).
222
Feijen: Revision of Eurydiopsis
helsdingeni sp. n. Type locality: Java, Indonesia.
Holotype in rmnh, 2 paratypes in irsnb, 1
para type in mluh.
Ref.: van der Wulp 1897: 189 (as 'Diopsis
anularis Macquart); Frey 1928: 70 (as 'Diopsis
anularis Macquart) .
saraivakensis sp. n.
Type locality: Sarawak, Malaysia. Holotype in
rmnh, 2 paratypes in bpbm.
'subnitidd; (Bigot 1881: 373, lapsus for subnotata
Westwood).
subnotata'Westwood, 1848, 37, pi. 18, fig. 2 (Diopsis).
Type locality: Philippines.
Ref.: Frey 1928: 71 (in part, others now included
in glabrostylus sp. n.). Other records from India
(Assam), Burma, Indonesia (Sumatra, Celebes)
and New Guinea (Irian Jaya) are probably all
misidentifications. Type ( 9 ) in bmnh.
Distribution
So far the various Eurydiopsis species were all lumped
under the name E. subnotata. This name was recorded
from the Philippines (Westwood 1848: 37, Bezzi 1913:
328, Frey 1928: 71), Assam in India (Brunetti 1907:
164), Myanmar (Steyskal 1975: 33), Celebes, Java and
Sumatra in Indonesia (Walker 1 861 : 263, van der Wulp
1896: 171, 1897: 192, de Meijere 1924: 60), mainland
Malaysia (Tan 1965: 14, 1966: 133, 1967: 36;
Burkhardt & de la Motte 1983: 99, 1985: 204) and
New Guinea (Kertész 1899: 183). However, Eurydiopsis
subnotata is now considered as a species which only oc-
curs in the south-eastern islands of the Philippines. As a
second species from the Philippines, Eurydiopsis
glabrostylus is now recognized. This species occurs in the
north/north-western islands of the Philippines. Eurydiop-
sis argentifera, formerly placed in synonymy with E sub-
notata, is the most widely distributed species, occurring
in Sulawesi (Celebes), Java, Sumatra and mainland
Malaysia. Male genitalia of the various locations were
compared (cf. figs. 1-3), but did not warrant the erection
of more species. Given the distribution of E. argentifera,
the possibility cannot be excluded that it also occurs in
Borneo, as the second species of that island, besides the
indigenous Eurydiopsis saraivakensis sp. n. The aberrant
Eurydiopsis helsdingeni is the second species from Java,
while Eurydiopsis brevispinus occurs in Laos and Myan-
mar. The Eurydiopsis recorded from Assam and from
New Guinea are likely to represent additional unde-
scribed species. Unfortunately, no material from these
locations could be traced.
Phylogeny
Feijen (1989) discussed our present knowledge of
the intrafamiliar relationships of the Diopsidae. For
the moment the Diopsidae are divided into the sub-
families Sphyracephalinae and Diopsinae. The Sphy-
racephalinae are likely to represent a monophyletic
group, but a proper groundplan with synapomorphic
conditions cannot yet be presented. It remains to be
seen whether the Diopsinae form a monophyletic
group, and, as such, the sistergroup of the Sphyra-
cephalinae. Feijen (1989) indicated a few possible
groups of genera within the Diopsinae, but Eurydiop-
sis belongs to the genera of which the position within
the Diopsinae remains unclear. In the key presented
by Feijen (1989) Eurydiopsis keys out next to Diopsis,
but this does not necessarily give an indication of a
closer relationship.
As important plesiomorphic characters of Eury-
diopsis, the following conditions can be given: absence
of sexual dimorphism in length of eye stalks and
shape of front femora, absence of facial teeth (or pres-
ence of very small rounded teeth) and presence of a
distinct seam between terga 2 and 3 in the syntergum.
A distinct seam between terga 2 and 3 in the synter-
gum further only occurs in Diasemopsis and Sinodiop-
sis, while in some species of the Diopsis indica com-
plex a vague seam occurs.
As important apomorphic characters the absence of
thoracic bristles and the very short, band-like male
sternum 7+8 (fig. 26) can be mentioned. The only
other genus in which no thoracic bristles occur is
Diopsis. A reduced sternum 7+8 is only found in
Sphyr acephala, Cladodiopsis, Eurydiopsis and the
Diopsis indica complex.
Intriguing is the occurrence of similar (subapical,
central and proximal rows of irrorations) wing pat-
terns in the genera Diopsina, Sinodiopsis, Eurydiopsis,
Teleopsis, Cyrtodiopsis and Eurydiopsis S. & V. nee
Frey. Feijen (1989) still assumed that these wing pat-
terns did not indicate a common origin. However,
the possibility cannot be excluded. Based on the com-
mon character of a very narrow male sternum 7+8, a
closer relationship between Eurydiopsis and the Diop-
sis indica complex can also not yet be ruled out. It is
doubtful that the indica complex will remain in a re-
vised Diopsis genus.
Within Eurydiopsis, the subnotata complex and E.
helsdingeni can be considered as sistergroups. As syn-
apomorphic condition for the subnotata complex the
slender to moderately incrassate front femora can be
mentioned, while as apomorphic conditions for E.
helsdingeni the fusion of tergum 4 to the syntergum,
the glabrous wingbase and the divided male sternum 5
can be given. Within the subnotata group it is difficult
to indicate relationships. It does not seem unlikely that
E. glabrostylus and E. brevispinus are more closely relat-
ed. This assumption is based on similar character states
as the relative length of the scutellar spines, the ratio
length/width of the gonostyli and the percentage cov-
erage of the gonostyli with microtrichia (see table 1).
223
Tijdschrift voor Entomologie, volume i4i, 1998
Figs. 1-9. Latero-ventral view of gonostylus. — 1, Eurydiopsis argentifera, Indonesia, Sulawesi, nr. Bantimurung; 2, Eurydiop-
sis argentifera, Indonesia, Java; 3, Eurydiopsis argentifera, Malaysia, Malaya (bred); 4, Eurydiopsis brevispinus, paratype, Myan-
mar, Mt. Victoria; 5, Eurydiopsis brevispinus, paratype, Laos, Pau; 6, Eurydiopsis glabrostylus, paratype, Philippines, Aroroy; 7,
Eurydiopsis helsdingeni, paratype, Indonesia, Java; 8, Eurydiopsis sarawakensis, paratype, Malaysia, Sabah; 9, Eurydiopsis sub-
notata, Philippines, Surigao. Scale 0. 1 mm.
Biology
Our knowledge of the biology of Eurydiopsis is
mainly based on the studies of Eurydiopsis argentifera
(as subnotata) by Burkhardt & de la Motte (1983: 99,
1985: 204), while a few observations were made by
Tan (1965: 14, 1967: 36).
Among the Diopsidae the Eurydiopsis species are
exceptional because of their size and weight. The
Philippine Eurydiopsis are the largest (length of body)
and heaviest diopsids known. However, there is noth-
ing unusual about their eye stalks, which are, relative-
ly, among the smaller ones in the Diopsidae. The ab-
sence of sexual dimorphism with regards to eye span
and shape of femora is found in all Eurydiopsis.
Burkhardt & de la Motte (1985: 204) also recorded
Eurydiopsis argentifera (as subnotata) as a homomor-
phic species with a sex ratio of freshly emerged flies
close to a 1 : 1 ratio.
Another remarkable feature of Eurydiopsis is their
unusual life span. Burkhardt & de la Motte (1983:
99) mentioned a life span of almost one-and-a-half
year for the Malayan Eurydiopsis argentifera.
Differential and quantitative characters
Measurements. - Within Eurydiopsis the two Phi-
lippine species, E. glabrostylus and subnotata are the
large species, argentifera and sarawakensis are medi-
um-sized, while helsdingeni and brevispinus are small.
The various quantitative data for the species can be
compared in table 1. The average relative eye span
(ratio eye span/body length) is very uniform among
the species concerned and varies only between 66%
and 71%. The categories used for description of the
relative eye span are given in Feijen (1998). Hardly
any difference occurs in the relative eye spans of fe-
males and males, although the relative eye span is for
all species marginally longer in the males. More inter-
specific variation occurs in the average relative length
of the scutellar spines (ratio scutellar spines/body
length). Large spines occur in E. argentifera, sarawak-
ensis and subnotata, with ratios varying from 24% to
26%. Smaller spines occur in E. brevispinus and
glabrostylus (ratios varying from 19% to 20%), while
the spines are with 15% smallest in E. helsdingeni.
Head. - Small, rounded facial teeth occur in E. ar-
gentifera and subnotata. In the other four species the
224
Feijen: Revision of Eurydiopsis
Table 1 . Measurements (in mm) and other quantitative characters for the six recognized Eurydiopsis species.
Character
Eurydiopsis
helsdingeni
Eurydiopsis subnotata complex
argentifera
brevispinus
glabrostylus sarawakensis
subnotata
9
3
?
3
9
S
9
3
3
9
3
Length of body
8.2
7.8
9.7
9.3
8.5
8.6
10.7
10.2
9.4 11
9.7
Eye span
5.4
5.7
6.6
6.4
5.4
5.7
6.9
7
6.6
7.5
6.9
Ratio eye span/
body length
0.66
0.73
0.68
0.71
0.64
0.66
0.66
0.69
0.7
0.68
0.71
Length of wing
5.4
5.3
7.1
6.7
6.6
6.4
7.2
7.1
7 7.9
6.8
Length of sc. spines
1.27
-
2.31
2.26
1.71
1.71
2.01
2.05
2.46 ; 2.73
2.41
Ratio scsp/body
0.15
-
0.24
0.24
0.2
0.2
0.19
0.2
0.26 0.25
0.25
Ratio 1/w of femur 1
3.6
3.6
5.1
5.1
5.1
4.7
5.4
5.5
5.1 5.3
5.3
Outer tub. of femur 1
18.7
19
20.6
20.3
18.7
19
19.3
21.8
20.2 22.5
22.5
Inner tub. of femur 1
22.3
24
23.1
23.2
22.8
23.5
22.1
24.3
22.5 26
24.7
Ratio 1/w of cerei
2.5
1.6
1.8
3.4
1.7
5
1.7
3.6
3.1 1.9
7.6
Ratio 1/w of gonostyli
10
1.5-1.6
2.4-2.7
2.7
1.7
2.3
% coverage of gono-
styli with microtrichia
<5
40-50
<5
<5
30
30
Flies examined
3
1
8
5
2
3
5
5
3
3
9
facial corners are rounded. Small differences occur in
the structural pattern and the pollinosity pattern of
the frons. The IVB is minuscule in all species, except
for E. sarawakensis, in which it is somewhat longer.
Wing. - In E. helsdingeni a large area of apical in-
fuscation occurs, in glabrostylus a small area of infus-
cation occurs at the apex, while in the other four
species no apical infuscation occurs at the tip. The
distal anterior hyaline spot is small in E. subnotata,
medium-sized in helsdingeni and large in the other
four species. This spot continues well into the first
posterior cell in E. argentifera, brevispinus, glabrostylus
and helsdingeni, while in sarawakensis and subnotata
the spot does not cross the third vein. The proximal
anterior hyaline spot and the wingbase are glabrous in
E. helsdingeni, while in the other five species the wings
are almost uniformly pollinose with only in the basal
region some tiny glabrous spots.
Legs. - The front femur is incrassate in E. helsdin-
geni (ratio length/width 3.6) and slightly incrassate to
slender (ratios varying between 4.7 and 5.5) in the
species of the subnotata complex. No sexual dimor-
phism could be found in this respect. In one species
(E. brevispinus) the femur is even slightly more incras-
sate in the male, while normally in dimorphic diop-
sids the male has more slender femora. The categories
used for description of the ratio length/width of the
front femur are given in Feijen (1998). The numbers
of tubercles in the two rows on the front femora can-
not serve as differential characters within Eurydiopsis
(table 1).
Preabdomen. - In E. helsdingeni the syntergum in-
cludes the first four terga, while in the species of the
subnotata complex only the first three terga are in-
cluded. According to Feijen (1989: 109) the only
other diopsid with a syntergum consisting of terga
1+2+3+4 is Cobiopsis latifascia (Brunetti 1928: 592).
Minor specific differences occur in the patterns of the
densely white pollinose spots.
Female postabdomen. - Tergum 8 is divided into
two sclerites in E. helsdingeni and glabrostylus, where-
as in the other species it is a single plate, with a medi-
al constriction in some species. Sternum 8 is strongly
constricted in E. helsdingeni, less constricted in argen-
tifera and brevispinus and not constricted in glabrosty-
lus and subnotata. The cerei are rather broad in E.
helsdingeni (ratio length/width 2.5) and broad in the
other species (ratios varying from 1.7-1.9. The fol-
lowing categories are used for description of both fe-
male and male cerei: broad if the ratio length/width
(L/W) is SS2.0, rather broad m case of 2.0<L/W«£3.0,
rather elongate for 3.0<L/W^4.0, elongate for
4.0<L/W=S5.0, and very elongate for 5.0<L/W. The
subanal plate is more or less pentagonal in E. helsdin-
geni, argentifera and subnotata, triangular in brevispi-
nus and somewhat rectangular with rounded posteri-
or corners in glabrostylus.
Male postabdomen. - Sternum 5 is divided in E.
helsdingeni and a single plate in the other species. The
ratio length/width of the gonostyli is a major differ-
ential character and varies from 10 in £ helsdingeni,
via 2.3-2.7 in brevispinus, glabrostylus and subnotata,
225
Tijdschrift voor Entomologie, volume hi, 1998
Figs. 10-21. Eurydiopsis argentifera. — 10, 9 , head in anterior view; 11, 3 , head in anterior view; 12, c?,wing; 13, S .dorsal
view of abdomen; 14, 9, ventral view of abdomen; 15, dorsal view of 9 terga 8, 10 and cerei; 16, ventral view of subanal
plate; 17, spermathecae; 18, genital ring; 19, posterior view of periandrium with gonostyli and cerei; 20, lateral view of phal-
lapodeme; 21, ejaculatory apodeme and sac. Scales: 1 mm (10-14), 0.1 mm (15-21). - Figs. 10, 15-21, Malaysia, Malaya
(bred), 13, Indonesia, Sulawesi, nr. Bantimurung.
226
Feijen: Revision of Eurydiopsis
to 1.5-1.7 in argentifera and sarawakensis (table 1).
The gonostyli are slightly constricted in E. brevispinus
and glabrostylus. In E. helsdingeni, brevispinus and
glabrostylus the gonostyli are largely glabrous with
only a few microtrichia apically, in sarawakensis and
subnotata the apical third of the outer side is covered
with microtrichia, whereas in argentifera almost the
whole apical half is pollinose. The ratio length/width
of the cerei varies from 1.6 to 7.6. However, the
shape of the cerei is rather irregular, making this char-
acter less useful.
Key to the species
1. Syntergum 1+2+3, base of wing pollinose, front
femur moderately incrassate to slender (ratio 1/w
> 4.5), S sternum 5 a single sclerite, ratio 1/w of
gonostylus < 3 (subnotata complex) 2
- Syntergum 1+2+3+4, base of wing glabrous,
front femur incrassate (ratio 1/w <4), S sternum
5 divided, ratio 1/w of gonostylus ±10, Java. . . .
helsdingeni
2. Small facial teeth present 3
- Facial teeth absent 4
3. Depression in front of ocellar tubercle, frons
pollinose, small distal anterior wingspot not ex-
tending into first posterior cell, 9 sternum 8 me-
dially not constricted, ratio 1/w of gonostyli ± 2.3,
Philippines subnotata
- Elevation in front of ocellar tubercle, frons glossy,
large distal anterior wingspot extending into first
posterior cell, 9 sternum 8 medially constricted,
ratio 1/w of gonostyli ± 1.5-1.6, Indonesia and
Malaya argentifera
4. Distal anterior wingspot extending into first pos-
terior cell, rvB a minuscule wart, scutellar spines
medium-sized (19-20% of body length), ratio 1/w
of gonostyli ± 2.4-2.7, gonostyli largely glabrous
with apically a few microtrichia 5
- Distal anterior wingspot not extending into first
posterior cell, rvB a small bristle, scutellar spines
large (± 26% of body length), ratio 1/w of gono-
styli ± 1 .7, apical third of outer side of gonostyli
pollinose, Borneo sarawakensis sp. n.
5. Wing apex slightly infuscated, 9 tergum 8 divid-
ed, subanal plate apically rounded, gonostyli
slightly constricted, S cercus with ratio 1/w ± 3.6,
Philippines glabrostylus sp. n.
- Wing apex hyaline, 9 tergum 8 a single sclerite,
subanal plate triangular, gonostyli not constrict-
ed, ô cercus with ratio 1/w ± 5.0, Laos and
Myanmar brevispinus sp. n.
Eurydiopsis argentifera (Bigot, 1874)
(figs. 1-3, 10-21)
Diopsis argentifera Bigot, 1874: 1 12. Two syntypes (as 'co-
types'), Celebes (Sulawesi), Indonesia (umo). [Examined]
Diopsis subnotata; Walker 1861: 263 (wrong identification);
Bigot 1881: 372 (senior synonym of argentifera, quoting
Osten Sacken 1882: 237); Osten Sacken 1882: 237 (se-
nior synonym of argentifera); van der Wulp 1896: 171,
1897: 192; de Meijere 1924: 60; Shillito 1971: 290-291:
Tan 1965: 14, 1966: 133, 1967: 36; Burkhardt & de la
Motte 1983: 99, 1985: 204.
Further material. - 19, le?, Indonesia, Sulawesi, nr.
Bantimurung, Pattunuang Asue, 17-20. iv. 1991, c60m, C.
v. Achterberg (rmnh); là, Indonesia, Sulawesi, Watuwila
Sulawesi SW 08, 16.X.1989, Krikken & Blom (rmnh); 1 S,
Indonesia, Java, Batoerraden, G. Slamat., 17. vii. 1927, F.C.
Drescher (zma); 19, Indonesia, Sumatra, Serapa Kur.,
vii. 191 5, Edw. Jacobson (zma); 1 S , Indonesia, Java, Blume
(rmnh); 1 9, Indonesia, W. Java, Tjidaon, Udjong Kulon,
5.XÜ.1958, A.N.R. Wegner (rmnh); 29, 4 e?, Malaya,
Malaysia 1988, bred by Burkhardt & de la Motte (rmnh).
Description
Measurements. - (The measurements of the bred
Malaysia flies were slightly smaller than those of wild
material) Length of body in 9 9.7 mm + se 0.2
(range 9.1-10.5) and in 6 9.3 mm ± 0.3 (range 7.4-
10.7), eye span in 9 6.6 mm ±0.1 (range 6.4-7.0)
and in cT 6.4 mm + 0.2 (range 5.2-7.0), length of
wing in 9 7. 1 mm ±0.1 (range 6.9-7.3) and in cT 6.7
mm ±0.1 (range 6.0-7.3), length of scutellar spine in
9 2.31 mm ± 0.10 (range 1.83-2.57) and in S 2.42
mm + 0.06 (range 2.05-2.54).
Head. - Central part black, thinly pollinose; ocel-
lar tubercle blackish; frons (fig. 10) with smooth ele-
vation in front of ocellar tubercle, lateral areas
smooth, a ridge around the frons; arcuate groove con-
colorous with rest of central part of head; face black-
ish, smooth and thinly pollinose, with ridge parallel
to and just below arcuate groove, face somewhat
bulging centtally, a few pale hairs, small blunt facial
teeth present (figs. 10-11); eye span very small in fe-
male (32% smaller than the length of body) and very
small in male (31% smaller than the length of body);
stalks dark brown, broad apical parts blackish, polli-
nose; rvB tiny, ove- medium-sized, slightly longer
than the diameter of the eye stalk.
Thorax. - Collar, scutum, scutellum, scutellar
spines, pleura and sterna uniformly blackish brown
pollinose; scutellar spines long (24% of body length),
more than 3X scutellum; almost straight, slightly
curved inward, diverging under an angle of 75-80°;
metapleural spines large, glossy, laterally directed;
very few sparse hairs on thorax.
Wing. - Apical eighth hyaline without infuscation
at the tip (fig. 12); three complete transverse bands;
227
Tijdschrift voor Entomologie, volume i4i, 1998
Figs. 22-33. Eurydiopsis brevispinus. — 22, S , head in anterior view; 23, c? , wing; 24, $ , dorsal view of abdomen; 25, 9 , ven-
tral view of abdomen; 26, 3 sternum 7+8; 27, dorsal view of terga 8, 10 and cerei; 28, ventral view of subanal plate; 29, sper-
mathecae; 30, genital ring; 31, posterior view of periandrium with gonostyli and cerei; 32, lateral view of phallapodeme; 33,
ejaculatory apodeme and sac. Scales: 1 mm (22-25), 0.1 mm (26-33). - Figs. 26, 31-33, holotype; 22, 23, 3 paratype, Laos,
Pao; 24, $ paratype, Laos, Pong King; 25, 27-30, 9 paratype, Myanmar.
228
Feijen: Revision of Eurydiopsis
preapical band darkest, extending from tip of posteri-
or crossvein to well beyond the tip of the second vein,
proximal edge and apical edge straight; central band
extending from base of submarginal cell to posterior
crossvein, rather vague, darker around anterior cross-
vein; basal band very vague and irregular hardly
reaching the anterior margin and constricted in anal
cell, extending from base of third posterior cell to tip
of anal cell; preapical band and central band broadly
connected in posterior half of first posterior cell, cen-
tral band and basal band connected in discal cell; ex-
cept for hyaline base five hyaline spots, one from tip
of costal cell extending to fourth vein, one from tip of
anal cell to almost the wing margin, one large and dis-
tinct one extending from anterior wing margin to
well into the first posterior cell, one basally in second
posterior cell and one occupying the apical eighth of
the wing; wing almost uniformly covered by mi-
crotrichia, only glabrous sections in base of costal cell
and basal half of anal cell.
Legs. - Front leg brown, with blackish brown coxa,
tibia and metatarsus, paler other tarsi, short black
stripe apically on femur, coxa 1 pollinose, pollinosity
on inner and outer side of femur 1 ; mid leg brown
with whitish basal 2/5 of femur; hind leg brown with
pale basal eighth of femur; femur 1 slender in 9 and
Ô (ratio of length/ width in both 9 and 6 5.1 ± se
0.1, range 4.8-5.3); tubercles on distal three-quarters,
inner row in 9 with 23.1 tubercles ±1.1 (range 18-
27) and in â with 23.2 tubercles ±1.0 (range 16-27),
outer row in 9 with 20.6 tubercles ±1.0 (range 17-
26) and in 6 with 20.3 tubercles ± 0.8 (range 17-24).
Preabdomen. - Dorsally blackish brown, pollinose,
base more whitish pollinose, terga 2 and 3 anterolat-
erally with whitish pollinose spots; tip (centre and
apical edge of tergum 4, and tergum 5) whitish polli-
nose (fig. 13 ): syntergum consisting of terga 1, 2 and
3, seam between terga 2 and 3 distinct; sternum 1
dark brown, other sterna brown with whitish polli-
nose apical bands, pollinose.
Female postabdomen. - Deflexed, terga 6 and 7 rec-
tangular plates (fig. 14: the female abdomens are in
these types of figures presented in a see through way, so
that the terga are visible behind the sterna); tergum 8
(fig. 1 5) represented by a single plate, posteriorly in a
V-shaped way constricted medially, basal half of ter-
gum 8 glabrous except medially; tergum 1 0 with three
pair of hairs: cerei broad, ratio of length/width 1.8,
covered with microtrichia and a number of hairs (fig.
15); sterna 5 and 6 single rectangular sclerites (fig. 14);
sternum 7 strongly constricted medially, almost cut in
two sclerites; sternum 8 a single, somewhat bean-
shaped sclerite, slightly constricted medially; subanal
plate (figs. 15-16) pentagonal, posteriorly nine pairs of
hairs; spermathecae (fig. 17) rounded and smooth:
genital ring (fig. 1 8) rounded.
Male postabdomen. - Terga 5 and 6 single plates;
sterna 4 and 5 single rectangular sclerites; sternum 6
represented by two small sclerites; sternum 7+8 a very
short, band-like sclerite; spiracles 5 and 6 in mem-
brane, spiracle 7 located rather dorsally, anteriorly of
sternum 7+8; periandrium (fig. 19) rounded, with
about 17 pairs of hairs, covered with microtrichia;
gonostyli rounded to rectangular in lateral view (figs.
1-3), ratio length/width 1.5-1.6, in posterior view
apically rounded, apical half on outer side covered
with microtrichia, on inner side glabrous, on outer
side some short hairs on apical third; gonostyli inter-
connected via thin processus longi; cerei rather elon-
gate, ratio length/width 3.4, broadest preapically,
covered with microtrichia and hairs; phallapodeme
(fig. 20) solidly built, not broadening anteriorly, an-
terior arm 1.5 times as long as posterior arm; ejacula-
tory apodeme (fig. 21) broadening anteriorly.
Diagnosis
Eurydiopsis argentifera belongs to the subnotata
complex and can be recognized by the wing pattern
(no infuscation at the tip, large distal anterior spot ex-
tending into first posterior cell), almost uniform dis-
tribution of microtrichia on the wing, tiny ivb, medi-
um-sized OVB, presence of facial teeth, slender front
femora, large scutellar spines (24% of body length),
syntergum 1+2+3, posteromedially constricted fe-
male tergum 8, medially strongly constricted female
sternum 7, somewhat bean-shaped female sternum 8,
broad female cerei, pentagonal subanal plate, single
male sternum 5, rounded to rectangular gonostyli
with a ratio length/width of 1.5-1.6, distal half of
outer side of gonostyli covered with microtrichia,
rather elongate male cerei with a ratio length/width of
3.4 and phallapodeme with anterior arm 1.5 times as
long as the posterior arm.
Eurydiopsis brevispinus sp. n.
(figs. 4-5, 22-30)
Type material. - S Holotype, 1 9 paratype, Myan-
mar (Burma), Mt. Victoria, Chinhills, 1000m,
iii.1938, leg. G. Heinrich (bmnh); 19 paratype,
Myanmar (Burma), S. Shan States, road 40 km E. of
Taunggyi, 1500m, 2-25.X.1934, Malaise (uzmh); 1 9
paratype, Laos, Pong King, 13. iv. 1918, R. Vitalis de
Salvaza (uzmh); 1 S paratype, Laos, Pau, 8 km w.,
18.iii.1920 (uzmh).
Description
Measurements. - Length of body in 9 8.5 mm ±
se 0.1 (range 8.3-8.6) and in S 8.6 mm (range 8.3-
8.8), eye span in 9 5.4 mm + 0.1 (range 5.3-5.5) and
in â 5.7 mm (range 5.6-5.7), length of wing in 9 6.6
mm ±0.1 (range 6.5-6.8) and in S 6.4 mm (range
229
Tijdschrift voor Entomologie, volume hi, 1998
6.1-6.6), length of scutellar spine in 9 1.71 mm ±
0.05 (range 1.61-1.77) and in 6 1.71 mm (range
1.64-1.77).
Head. - Central part blackish, thinly pollinose,
ocellar tubercle black; frons (fig. 22) somewhat vari-
able with slight, smooth elevation in front of ocellar
tubercle, slight depression below elevation, lateral
areas smooth, with circular ridge around the frons; ar-
cuate groove concolorous; face with ridge parallel to
and just below arcuate groove, face somewhat bulging
centrally, a few pale hairs, facial corners rounded; eye
span very small in female (36% smaller than the
length of body) and very small in male (34% smaller
than the length of body) ; stalks brown, broad apical
parts blackish, pollinose; ivb minuscule and wart-
like; OVB medium-sized, somewhat longer than the
diameter of the eye stalk.
Thorax. - Collar, scutum, scutellum, scutellar
spines, pleura and sterna uniformly blackish brown
pollinose; scutellar spines medium-sized (20% of
body length), almost 3X scutellum; almost straight,
slightly curved inward, diverging under an angle of
more than 80°; metapleural spines large glossy, later-
ally directed; some hairs on thorax.
Wing. - Apical eighth hyaline without infuscation
at the tip (fig. 23); three complete transverse bands,
preapical band darkest and with darker anterior half,
extending from tip of posterior crossvein to well be-
yond the tip of the second vein, proximal edge curved
and apical edge straight; central band extending from
base of submarginal cell to posterior crossvein, rather
vague, darker around anterior crossvein; basal band
very vague and irregular hardly reaching the anterior
margin and constricted in anal cell, extending from
base of third posterior cell to tip of anal cell; preapical
band and central band broadly connected around
fourth vein, central band and basal band connected
around fifth vein; except for hyaline base five hyaline
spots, one from tip of costal cell extending into discal
cell, one from tip of anal cell to wing margin, one
large and distinct one extending from anterior wing
margin to well into the first posterior cell, one basally
in second posterior cell and one occupying the apical
eighth of the wing; wing almost uniformly covered by
microtrichia, glabrous sections include most of costal
cell and basal parts of second basal cell and anal cell.
Legs. - Front leg brown, with darker coxa, tibia and
metatarsus, paler other tarsi, coxa densely pollinose,
remainder of front leg thinly pollinose; mid leg and
hind leg brown, apices of femur darker; femur 1 slen-
der in female (ratio of length/width 5.1 ± se 0.1, range
4.9-5.2) and moderately incrassate in male (ratio of
length/width 4.7, range 4.6-4.8); tubercles on distal
two-thirds, inner row in 9 with 22.8 tubercles ±0.5
(range 21-25) and in 6 with 23.5 tubercles (range 23-
24), outer row in 9 with 18.7 tubercles ±0.8 (range
16-21) and in 6 with 19.0 tubercles (range 19).
Preabdomen. - Dorsally blackish brown, pollinose,
base more whitish pollinose, terga 2 and 3 anterolat-
erally with whitish pollinose spots; tip (centre and
apical edge of tergum 4, and tergum 5) whitish polli-
nose (fig. 24): syn tergum consisting of terga 1, 2 and
3, seam between terga 2 and 3 distinct; sternum 1
dark brown, other sterna brown with whitish polli-
nose apical bands, pollinose.
Female postabdomen. - Deflexed, terga 6 and 7
rectangular plates (fig. 25), tergum 8 (fig. 27) slightly
constricted medially, tergum 8 covered with mi-
crotrichia except in anterolateral corners; tergum 10
with three pair of hairs: cerei broad, ratio of
length/width 1.7, covered with microtrichia and a
number of hairs (fig. 27); sterna 5 and 6 single rec-
tangular sclerites (fig. 25); sternum 7 also rectangular
but concave posteriorly; sternum 8 a single V-shaped
sclerite; spiracle 7 in membrane; subanal plate (figs.
27-28) triangular to kidney-shaped, posteriorly nine
pairs of hairs; spermathecae (fig. 29) rounded and
smooth: genital ring (fig. 30) rounded.
Male postabdomen. - Terga 5 and 6 single plates;
sterna 4 and 5 single rectangular sclerites; sternum 6
represented by two small triangular sclerites; sternum
7+8 a very short, band-like sclerite (fig. 26); spiracles
5 and 6 in membrane, spiracle 7 located rather dor-
sally, anteriorly of sternum 7+8 (fig. 26); periandrium
(fig. 3 1 ) rounded, with about 23 pairs of relatively
long hairs, covered with microtrichia; gonostyli ob-
long in lateral view with rounded apical corners (figs.
4-5), ratio length/width 2.4-2.7, very slightly con-
stricted in the middle, in posterior view apically
rounded, only the very apex on inner and outer side
with a few microtrichia, on outer side relatively long
hairs on apical half; gonostyli interconnected via thin
processus longi; cerei elongate, ratio length/width
5.0, broadest preapically, covered with microtrichia
and hairs; phallapodeme (fig. 32) rather slender, not
broadening anteriorly, anterior arm slightly longer
than posterior arm; ejaculatory apodeme (fig. 33)
broadening anteriorly.
Diagnosis
Eurydiopsis brevispinus belongs to the subnotata com-
plex and can be recognized by its wing pattern (no in-
fuscation at the tip, large distal anterior spot extending
into first posterior cell), almost uniform distribution of
microtrichia on the wing, minuscule ivb, medium-
sized OVB, absence of facial teeth, moderately incrassate
to slender front femora, medium-sized scutellar spines
(20% of body-length), syn tergum 1+2+3, rectangular
(medially slighdy constricted) female tergum 8, almost
rectangular sternum 7, V-shaped female sternum 8,
broad female cerei, triangular subanal plate, single male
sternum 5, oblong gonostyli with a ratio length/width
230
Feijen: Revision of Eurydiopsis
Figs. 34-43. Eurydiopsis glabrostylus. — 34, ? head in anterior view; 35, 9 wing; 36, S dorsal view of abdomen; 37, 9 ven-
tral view of abdomen; 38, dorsal view of 9 terga 8, 10 and cerei; 39, ventral view of subanal plate; 40, spermathecae; 41, pos-
terior view of periandrium with gonostyli and cerei; 42, lateral view of phallapodeme; 43, ejaculatory apodeme and sac. Scales:
1 mm (34-37), 0.1 mm (38-43). -All paratypes, Philippines, Aroroy.
231
Tijdschrift voor Entomologie, volume i4i, 1998
of 2.4-2.7, glabrous gonostyli with only apically a few
microtrichia, elongate male cerei with a ratio length/
width of 5.0 and phallapodeme with anterior arm only
slightly longer than posterior arm.
Eurydiopsis glabrostylus sp. n.
(figs. 6, 34-43)
Type material. - S holotype, 1 S paratype, Philip-
pines, Masbate, Aroroy, x.1917 (uzmh); 1$
paratype, Philippines, Masbate, Aroroy, viii.1918
(uzmh); 1 9 paratype, ló* paratype, Philippines, Lu-
zon, Los Banos, iv.1914 (uzmh); 2 9 paratypes, lo*
paratype, Philippines, Polillo, 18.viii.1915 (uzmh);
1 9 paratype, 1 S paratype, Philippines, Mindoro,
Subaan, i. 191 6 (uzmh).
Description
Measurements. - Length of body in 9 10.7 mm ±
se 0.6 (range 8.5-12.4) and in â 10.2 mm ± 0.4
(range 8.8-1 1.2), eye span in 9 6.9 mm ± 0.4 (range
5.5-8.1) and in 3 7.0 mm ± 0.3 (range 5.7-7.4),
length of wing in 9 7.2 mm ± 0.4 (range 6.0-8.3)
and in S 7.1 mm ± 0.3 (range 6.1-7.5), length of
saltellar spine in 9 2.01 mm ± 0.08 (range 1.74-
2.23) and in 6 2.05 mm ± 0.05 (range 1.86-2.14).
Head. - Central part blackish brown, pollinose;
ocellar tubercle blackish; frons (fig. 34) with smooth
elevation in front of ocellar tubercle, lateral areas
smooth, or provided with vague radiating grooves lat-
erally and anteriorly, a vague circular ridge around
the frons; arcuate groove concolorous with rest of
central part of head; face with ridge parallel to and
just below arcuate groove, face somewhat bulging
centrally, a few pale hairs, facial corners angular, but
no distinct facial teeth; eye span very small in female
(34% smaller than the length of body) and very small
in male (31% smaller than the length of body); stalks
dark brown, broad apical parts blackish, pollinose;
IVB a minuscule wart, ovb small, 1.5X the diameter
of the eye stalk.
Thorax. - Collar, scutum, scutellum, scutellar
spines, pleura and sterna uniformly blackish brown
pollinose; scutellar spines medium-sized (19% of
body length), about 3X scutellum; slightly curved in-
ward, diverging under an angle of 75°; metapleural
spines large glossy, laterally directed; some hairs on
thorax.
Wing. - Apical tenth hyaline with some distinct in-
fuscation at the tip (fig. 35); three complete trans-
verse bands, preapical band darkest, extending from
tip of posterior crossvein to well beyond the tip of the
second vein, convex on proximal edge and convex on
apical edge; central band extending from base of sub-
marginal cell to posterior crossvein, rather vague,
darker around anterior crossvein and in first posterior
cell; basal band very vague and irregular hardly reach-
ing the anterior margin, extending from base of third
posterior cell to tip of anal cell; preapical band and
central band broadly connected around fourth vein,
central band and basal band connected in and around
discal cell; except for hyaline base five hyaline spots,
one from tip of costal cell extending to fourth vein,
one from tip of anal cell to almost the wing margin,
one large and distinct one extending from anterior
wing margin to well into the first posterior cell, one
basally in second posterior cell and one occupying
most of the apical tenth of the wing; wing almost uni-
formly covered by microtrichia, only a tiny glabrous
section in base of costal cell.
Legs. - Front leg brown with blackish brown tibia,
coxa and metatarsus, paler other tarsi, densely polli-
nose coxa and thinly pollinose femur; mid leg and
hind leg brown with darker apical sections on femora
and slightly darker tibiae; femur 1 slender in 9 (ratio
of length/width 5.4 ± se 0.1, range 5.0-5.6) and slen-
der in â (ratio of length/width 5.5 ±0.1, range 5.2-
5.8); tubercles on distal three-quarters, inner row in
9 with 22.1 tubercles ±0.9 (range 19-26) and in 6
with 24.3 tubercles ±0.8 (range 22-27), outer row in
9 with 19.3 tubercles ± 0.6 (range 16-22) and in S
with 21.8 tubercles ± 1.0 (range 17-24).
Preabdomen. - Dorsally blackish brown, pollinose,
base more whitish pollinose, terga 2 and 3 anterolat-
erally with whitish pollinose spots; tip (including
most of tergum 4 and all of tergum 5) also whitish
pollinose (fig. 36): syntergum consisting of terga 1, 2
and 3, seam between terga 2 and 3 distinct; sternum
1 dark brown, other sterna brown with whitish polli-
nose apical bands, pollinose.
Female postabdomen. - Deflexed, terga 6 and 7
rectangular plates (fig. 37); tergum 8 (fig. 38) repre-
sented by two rectangular plates, separated medially,
basal half of tergum 8 glabrous except medially; ter-
gum 1 0 with four pairs of hairs, cerei broad, ratio of
length/width 1.7, covered with microtrichia and a
number of hairs (fig. 38); sterna 5 and 6 single rec-
tangular scleri tes (fig 37); sternum 7 angular anterior-
ly and strongly concave posteriorly; sternum 8 a sin-
gle, somewhat bean-shaped sclerite; subanal plate
(figs. 38-39) somewhat rectangular with rounded
posterior corners, posteriorly nine pairs of hairs; sper-
mathecae (fig. 40) rounded and smooth.
Male postabdomen. - Terga 5 and 6 single plates;
sterna 4 and 5 single rectangular sclerites; sternum 6
represented by two small sclerites; sternum 7+8 a very
short, band-like sclerite; spiracles 5 and 6 in mem-
brane, spiracle 7 located rather dorsally, anteriorly of
sternum 7+8; periandrium (fig. 41) rounded, with
about 16 pairs of hairs, covered with microtrichia;
gonostyli oblong in lateral view (fig. 6), slightly but
distinctly constricted in the middle and with rounded
232
Feijen: Revision of Eurydiopsis
apical corners, ratio length/width 2.7, in posterior
view apically pointed, distally on outer side a few mi-
crotrichia, on inner side glabrous, on outer side some
short hairs on apical half; gonostyli interconnected
via thin processus longi; cerei rather elongate, ratio
length/width 3.6, broadest preapically, covered with
microtrichia and hairs; phaJlapodeme (fig. 42) solidly
built, broadening anteriorly, anterior arm just as long
as posterior arm; ejaculatory apodeme (fig. 43) broad-
ening anteriorly.
Diagnosis
Eurydiopsis glabrostylus belongs to the subnotata
complex and can be recognized by its wing pattern
(some infuscation at the tip, large distal anterior hya-
line spot extending into first posterior cell), almost
uniform distribution of microtrichia on the wing, mi-
nuscule rvB, small ovb, absence of facial teeth, slender
front femora, medium-sized scutellar spines (19% of
body length), syntergum 1+2+3, divided female ter-
gum 8, posteriorly strongly concave female sternum 7,
bean -shaped female sternum 8, broad female cerei, rec-
tangular subanal plate with rounded posterior corners,
single male sternum 5, oblong gonostyli slightly con-
stricted in the middle and with a ratio length/width of
2.7, glabrous gonostyli with only apically a few mi-
crotrichia, male cerei with a ratio length/width of 3.6
and phallapodeme with equal-sized arms.
Eurydiopsis hehdingeni sp. n.
(figs. 7, 44-55)
Diopsis circularise van der Wulp, 1897: 189 (nee circularis
Macquart).
Diopsis circularis; Frey, 1928: 70 (nee circularis Macquart).
Type material. - 9 Holotype, Indonesia, Java,
Blume, Coll. F.M. v. d. Wulp (rmnh); 1 2 , 1 <J para-
type, Indonesia, Java, Fruhstorfer (irsnb); 1 $ para-
type, Indonesia, Java, Batavia (= Djakarta), von
Roder (mluh). The first three specimens were identi-
fied by van der Wulp (1897) as 'Diopsis circularis'.
The second specimen was also by E. Brunetti identi-
fied as ' D. circularis'.
Etymology. - It is my pleasure to name this inter-
esting species after Dr. P. J. van Helsdingen, curator
of Diptera at the Leiden Museum. Dr. van Helsdin-
gen originates from the same island as Eurydiopsis
hehdingeni sp. n.
Description
Measurements. - Length of body in 9 8.2 mm
(range 7.8-8.6) and in ô 7.8 mm, eye span in 9 5.4
mm (range 5.2-5.7) and in cT 5.7 mm, length of wing
in 9 5.4 mm (range 5.2-5.7) and in o* 5.4 mm,
Fig. 44. Eurydiopsis helsdingeni. - 9 paratype, Indonesia,
Java, habitus in dorsal view. Scale: 1 mm.
length of scutellar spine in 9 1.27 mm (range 1.24-
1.30), scutellar spines broken in cT.
Head. — Central part uniformly brown (not black
as stated by van der Wulp 1897), thinly pollinose;
frons (figs. 44-45) with small rounded elevation in
front of ocellar tubercle, anteriorly and laterally of
ocellar tubercle some small depressions, lateral areas
smooth, a ridge around the frons; arcuate groove dark
brown; face with ridge parallel to and just below ar-
cuate groove, smooth, lower fifth somewhat de-
pressed, a few pale hairs, facial corners rounded, defi-
nitely without very small facial teeth as stated by van
der Wulp 1897; eye span very small in female (34%
smaller than the length of body) and very small in
male (27% smaller than the length of body); stalks
brown, broad apical parts blackish, pollinose, funicu-
lus pale brown; rvB just indicated as a minuscule
black wart (not absent as mentioned by van der
Wulp); ovb medium-sized, 1.5X the diameter of the
stalk.
Thorax. — Collar dark brown to blackish brown,
pollinose; scutum, scutellum and scutellar spines con-
colorous with collar, pollinose; pleura and sterna also
233
Tijdschrift voor Entomologie, volume mi, 1998
Figs. 45-55. Eurydiopsis helsdingeni. - 45, 9 , head in anterior view; 46, 9 , wing; 47, 9 , front leg, outer side; 48, 9 ventral
view of abdomen; 49, dorsal view of 9 terga 8, 10 and cerei; 50, ventral view of subanal plate; 51, spermathecae; 52, genital
ring; 53, posterior view of periandrium with gonostyli and cerei; 54, lateral view of phallapodeme and aedeagus; 55, ejacula-
tory apodeme and sac. Scales: 1 mm (45-48), 0.1 mm (49-55). - Figs. 48-52, holotype, Indonesia, Java; 45, 53-55, paratypes,
Indonesia, Java; 46, 47, 9 paratype, Indonesia, Java, Batavia.
234
Feijen: Revision of Eurydiopsis
concolorous and uniformly pollinose; scutellar spines
(fig. 44) relatively small (15% of body length), almost
2.5 X scutellum, pointed slightly upward, tip curving
slightly inward, diverging under an angle of 90°;
metapleural spines rather large, laterally directed; a
sparse distribution of fine hairs on thorax, especially
on the scutellar spines.
Wing. - Apex with large patch of infuscation (fig.
46); three complete transverse dark bands, the bands
together with the infuscated apex giving van der
Wulp's 'vier bruine dwarsbanden' (four brown cross-
bands); preapical band darkest and broadest, extend-
ing from tip of posterior crossvein to apically of tip of
second vein; central band extending from base of sub-
marginal cell to tip of fifth vein: basal band vague and
irregular, hardly reaching the anterior margin and
constricted in anal cell, extending from base of third
posterior cell to apex of anal cell; basal and central
band connected around fifth vein, central band and
subapical band connected around fourth vein; except
for hyaline base five hyaline spots, one from tip of
costal cell extending to just in discal cell, one central-
ly in third posterior cell, one extending from subapi-
cally in the marginal cell to halfway in the first poste-
rior cell, one basally in second posterior cell just
extending in discal cell and one hyaline subapical
crossband between dark subapical band and infuscat-
ed tip; the glabrous basal wing pans include most of
the subcostal cell, basal tip of marginal cell, basal half
of first basal cell, basal tip of second basal cell and
most of anal cell, the hyaline spot below the tip of the
subcostal cell is also glabrous. Van der Wulp indicat-
ed the differences with the wing pattern of Mac-
quart's circularis, but thought these due to inaccura-
cies and 'phantasie' of Macquart (1835). However,
Macquart's inaccuracy consisted only of mentioning
Java as a location of the African circularis. Macquart
made exactly the same error with his subfasciata (see
Feijen 1978).
Legs. — Front leg brown, coxa, tibia and metatarsus
dark brown, thinly pollinose; mid leg and hind leg
brown with darker tibiae, pollinose; femur 1 (fig. 47)
incrassate in $ (ratio of length/width 3.5, range 3.4-
3.5) and incrassate in 6 (ratio of length/width 3.6);
tubercles on distal three-quarters, inner row in 9 with
22.3 tubercles (range 22-23) and in cT with 24 tuber-
cles, outer row in ? with 18.7 tubercles (range 18-19)
and in â with 19 tubercles, outer row without gap.
Preabdomen. - Dorsally dark brown, uniformly
pollinose, tergum 3 (fig. 44) anterolaterally with
densely pollinose spots; basally strongly constricted,
posteriorly (tergum 4) rounded; syntergum consisting
of the terga 1, 2, 3 and 4, seam between terga 2 and 3
distinct, seam between terga 3 and 4 less distinct; ster-
na pale brown, sternum 1 darker; spiracle 1 in tergum
(fig. 48) other spiracles in membrane.
Female postabdomen. - Strongly deflexed under
syntergum 1-4 (figs. 44, 48); terga 6 and 7 rectangu-
lar plates (fig. 48); tergum 8 (fig. 49) represented by
two sclerites narrowly separated medially, sclerites
covered by microtrichia; tergum 10 with four pairs of
hairs, no long central hairs; cerei rather broad, ratio of
length/width 2.5, covered with microtrichia and a
number of hairs (fig. 49); sterna 5, 6 and 7 single rec-
tangular sclerites (fig. 48), sternum 6 more sclerotized
posteriorly; sternum 8 strongly constricted medially,
the two halves narrowly connected posteriorly; spira-
cle 7 in membrane; subanal plate (figs. 49-50) almost
triangular with rounded lateral corners, posteriorly
two pairs of long hairs and two pairs of short hairs,
the central pair of long hairs quite pronounced; sper-
mathecae (fig. 51) rounded, apically concave but this
may be due to an artefact, smooth; genital ring
rounded (fig. 52).
Male postabdomen. - Terga 5 and 6 rectangular
sclerites; sternum 4 a single rectangular sclerite; ster-
num 5 represented by a pair of L-shaped sclerites; ster-
num 6 represented by a pair of tiny sclerites: sternum
7+8 a short, band-like sclerite; spiracle 5 in mem-
brane, spiracle 6 just in membrane, spiracle 7 located
rather dorsally, anteriorly of sternum 7+8; periandri-
um (fig. 53) rounded, with about 22 pairs of hairs,
covered with microtrichia; gonostyli (fig. 7) long and
slender, ratio length/width about 10, glabrous with
apically some hairs and microtrichia, fused to perian-
drium but on cursory inspection apparently articulate;
gonostyli via thin processus longi connected to poste-
rior edge of periandrium and from there interconnect-
ed via thin processus; cerei broad, ratio of
length/width 1.6, covered with microtrichia and hairs,
apically some long hairs; phallapodeme (fig. 54) rather
slender, anterior arm slightly broadening anteriorly
and one third longer than posterior arm; ejaculatory
apodeme small, wedge-shaped (fig. 55).
Diagnosis
Eurydiopsis helsdingeni takes up an isolated position
in its genus, all other known species belonging to the
subnotata complex. It can be recognized by its wing
pattern (three dark crossbands and infuscated apex, five
hyaline spots), distribution of microtrichia on the wing
(glabrous base and glabrous basal anterior spot), mi-
nuscule IVB, medium-sized ovb, absence of facial teeth,
incrassate front femora, small scutellar spines (15% of
body length), syntergum 1+2+3+4 with distinct seam
between 2 and 3 and less distinct seam between 3 and
4, divided female tergum 8, rectangular female ster-
num 7, triangular subanal plate, rather broad female
cerei, divided male sternum 5, long, slender and almost
glabrous gonostyli with ratio length/width of 1 0, broad
male cerei and phallapodeme with anterior arm one
third longer than posterior arm.
235
Tijdschrift voor Entomologie, volume ui, 1998
Figs. 56-61. Eurydiopsis sarawakensis, male. - 56, head in anterior view; 57, wing; 58, dorsal view of abdomen; 59, poste-
rior view of periandrium with gonostyli and cerei; 60, ejaculatory apodeme and sac; 61, lateral view of phallapodeme. Scales:
1 mm (56-58 ), 0.1 mm (59-61). - Figs. 56, 58 holotype, 57 and 59-61, paratype, Malaysia, Sabah, Tenompok.
Eurydiopsis sarawakensis sp. n.
(figs. 8, 56-61)
Type material. - 6 Holotype, Malaysia, Sarawak,
Batu Niah, 29.xi.-27.xii. 1980, A. Harman (rmnh);
IS, Malaysia, Sabah (N. Borneo) or Sarawak: 1958-
1959, T.C. Maa (bpbm); 1<$, Malaysia, Sabah
(British Borneo), Tenompok, 1460m, Jesselton 30
mi. E., 10-19.ii.1959, T.C. Maa (bpbm).
Description
Measurements. - Length of body in S 9.4 mm ±
se 0.3 (range 8.8-9.9), eye span in S 6.6 mm ±0.1
(range 6.5-6.8), length of wing in o* 7.0 mm ± 0.1
(range 6.8-7.2), length of scutellar spine in â 2.46
mm ± 0.07 (range 2.33-2.54).
Head. - Central part dark brown, pollinose; ocellar
tubercle blackish; frons (fig. 56) with elevation in front
of ocellar tubercle and with small circular depression
in front of elevation, surrounded by V-shaped depres-
sion, lateral areas almost smooth, with some very
vague grooves, a circular groove around the frons; ar-
cuate groove concolo rous with central section of head;
face with ridge parallel to and just below arcuate
groove, face somewhat bulging centrally, a few pale
hairs, facial corners angular but without distinct facial
teeth; eye span very small in male (30% smaller than
the length of body); stalks brown, broad apical parts
blackish, pollinose; rvB small, not more than half the
length of the diameter of the eye stalk; ovb medium-
sized, somewhat longer than the diameter of the stalk.
Thorax. - Collar, scutum, scutellum, scutellar spines,
pleura and sterna uniformly blackish brown pollinose;
scutellar spines long (26% of body length), 3.5 X
scutellum; almost straight, very slightly curved inward,
diverging under an angle of 75°; metapleural spines
large glossy, laterally directed; some hairs on thorax.
Wing. - Apical eighth almost hyaline without in-
fuscation at the tip (fig. 57); three complete trans-
verse bands: preapical band darkest, extending from
tip of posterior crossvein to well beyond the tip of the
second vein, proximal edge slightly convex and apical
edge straight; central band extending from just before
base of submarginal cell to just past posterior cross-
vein, rather vague, darker around anterior crossvein
and around fifth vein; basal band very vague and ir-
regular hardly reaching the anterior margin and con-
stricted in anal cell, extending from base of third pos-
terior cell to tip of anal cell; preapical band and
central band broadly connected in whole of first pos-
terior cell and in anterior part of second posterior cell,
236
Feijen: Revision of Eurydiopsis
central band and basal band connected in whole of
discal cell; except for hyaline base five hyaline spots,
one just not reaching tip of costal cell and extending
to fourth vein, one from tip of anal cell to halfway the
wing margin, one distinct one extending from anteri-
or wing margin to third vein, one basally in second
posterior cell near wing margin and one occupying
the apical eighth of the wing; wing almost uniformly
covered by microtrichia, glabrous sections include
basal half of costal cell and basal parts of second basal
cell and anal cell.
Legs. - Front leg brown, blackish brown tibia, coxa
and metatarsus, paler other tarsi; coxa densely Polli-
nose, remainder of leg thinly pollinose, with more
dense pollinosity on sides of femur, mid leg and hind
leg brown with darker apical half on femur 2, two
dark bands on femur 3 and darker tibiae; femur 1
slender in male (ratio of length/width 5.1, range 5.0-
5.2); tubercles on distal two-thirds, inner row in S
with 22.5 tubercles ± se 0.8 (range 19-24), outer row
in cT with 20.2 tubercles ± 0.9 (range 18-23).
Preabdomen. - Dorsally blackish brown, pollinose,
base more whitish pollinose, terga 2 and 3 anterolat-
erally with whitish pollinose spots; tip (centre and
apical edge of tergum 4, and tergum 5) whitish polli-
nose (fig. 58): syntergum consisting of terga 1, 2 and
3, seam between terga 2 and 3 distinct; sternum 1
dark brown, other sterna brown with whitish polli-
nose apical bands, pollinose.
Male postabdomen. - Terga 5 and 6 single plates;
sterna 4 and 5 single rectangular sclerites; sternum 6
represented by two small kidney-shaped sclerites;
sternum 7+8 a very short, band-like sclerite; spiracles
5 and 6 in membrane, spiracle 7 located rather dor-
sally, anteriorly of sternum 7+8; periandrium (fig. 59)
rounded, with about 20 pairs of hairs, covered with
microtrichia; gonostyli rectangular in lateral view
with rounded apical corners, slightly constricted in
the middle (fig. 6), ratio length/width 1.7, in posteri-
or view apically pointed, on outer side the apical half
covered with microtrichia, inner side glabrous, on
outer side some hairs on apical half; gonostyli inter-
connected via thin processus longi; cerei rather elon-
gate, ratio length/width 3.1, broadest preapically,
covered with microtrichia and hairs; phallapodeme
(fig. 61) rather slender, not broadening anteriorly, an-
terior arm curving downward anteriorly and slightly
longer than posterior arm; ejaculatory apodeme (fig.
60) broadening anteriorly.
Diagnosis
Eurydiopsis sarawakensis belongs to the subnotata
complex and can be recognized by its wing pattern
(no infuscation at the tip, large distal anterior spot
not extending into first posterior cell), almost uni-
form distribution of microtrichia on the wing, small
IVB, medium-sized ovb, absence of facial teeth, slen-
der front femora, large scutellar spines (26% of body
length), syntergum 1+2+3, rectangular and in the
middle somewhat constricted gonostyli with a ratio
length/width of 1 .7, rather elongate male cerei with a
ratio length/width of 3.1 and phallapodeme with an-
terior arm slightly longer than posterior arm.
Eurydiopsis subnotata (Westwood, 1 848)
(figs. 9, 62-72)
Diopsis subnotata Westwood, 1848: 37, pi 18, fig 2. Type
( 9 ) in BMNH, carrying a label 'identified as the type by E.
E. Austen, from Philippine Is. Purchd fr. Mr. Wood
45.49'. [Examined]
Diopsis (Eurydiopsis) subnotata: Frey 1928: 71 (in part, the
remainder being glabrostylus sp. n.).
Eurydiopsis subnotata: Steyskal 1972: 11 (in part); Steyskal
1975: 33 (in pan); Feijen 1989: 62.
nee Diopsis argentifera Bigot, 1874: 112.
nee 'Diopsis subnotata'; van der Wulp 1896: 171, 1897: 192;
de Meijere 1924: 60, Tan 1965: 14, 1966: 133, 1967: 36;
Burkhard! & de la Motte 1983: 99, 1985: 204.
Further material. - 1 9, Philippines, Sororro, 17.V.1916
(uzmh); 19, Philippines, Samar, Catbalogan, iv.1915
(uzmh); 29, Philippines, Mindanao, Surigao, xi. 1915
(bmnh - Received from Dr. R. Frey -, one with label 'Diop-
sis subnotata Westw. Det. 1968, J.A. Tenorio'); 29, 2â ,
Philippines, Surigao, viii.1914 (uzmh); 19, Id, Philip-
pines, Mindanao, Surigao, xi.1915 (uzmh); 19, Philip-
pines, Surigao, viii.1916 (uzmh).
Description
Measurements. - Length of body in 9 1 1.0 mm ±
se 0.2 (range 10.1 -11.8) and in 6 9.7 mm ± 0.5
(range 8.8-10.5), eye span in 9 7.5 mm ±0.1 (range
7.3-7.7) and in S 6.9 mm ± 0.4 (range 6.1-7.4),
length of wing in ? 7.9 mm ±0.1 (range 7.4-8.2)
and in â 6.8 mm ± 0.3 (range 6.2-7.3) length of
scutellar spine in 9 2.73 mm ± 0.06 (range 2.39-
2.98) and in S 2.41 mm ± 0.05 (range 2.05-2.60).
Head. - Central part dark brown, pollinose; ocellar
tubercle blackish; frons (fig. 62) with ocellar tubercle
on elevation surrounded by sutures, small depression
in front of ocellar tubercle, lateral areas smooth or
provided with vague radiating grooves laterally and a
vague ridge around the frons; arcuate groove concol-
orous with rest of central part of head; face with ridge
parallel to just below arcuate groove, face somewhat
bulging centrally, a few pale hairs, facial corners with
distinct small facial teeth: eye span very small in fe-
male (32% smaller than the length of body) and very
small in male (29% smaller than the length of body);
stalks brown, broad apical parts blackish, pollinose;
funiculus paler brown, ivb minuscule; ovb medium-
sized, slightly longer than the diameter of the stalk.
Thorax. - Collar, scutum, scutellum, scutellar
spines, pleura and sterna uniformly blackish brown
237
Tijdschrift voor Entomologie, volume mi, 1998
Figs. 62-72. Eurydiopsis subnotata. - 62, ? , head in anterior view; 63, 5 , wing; 64, 9 , dorsal view of abdomen; 65, 9 , ven-
tral view of abdomen; 66, dorsal view of 9 terga 8, 10 and cerei; 67, ventral view of subanal plate; 68, spermathecae; 69, gen-
ital ring; 70, posterior view of periandrium with gonostyli and cerei; 71, lateral view of phallapodeme; 72, ejaculatory
apodeme and sac. Scales: 1 mm (62-65), 0.1 mm (66-72). - Figs. 62, 63, 65-72, Philippines, Mindanao, Surigao; 64, Philip-
pines, Sororro.
238
Feijen: Revision of Eurydiopsis
pollinose, scutellar spines long (25% of body length),
3.5 X scutellum; almost straight, slightly curved in-
ward, diverging under an angle of 75°; metapleural
spines large, glossy, laterally directed; few sparse hairs
on thorax.
Wing. - Apex hyaline with hardly any infuscation
(fig. 63) at the tip; three complete transverse bands,
preapical band darkest (especially anteriorly) and
slightly broader than central band, extending from tip
of posterior crossvein to well beyond the tip of the sec-
ond vein, apical edge of preapical band straight; cen-
tral band extending from base of submarginal cell to
posterior crossvein, rather vague, darker around ante-
rior crossvein; basal band very vague and irregular,
hardly reaching the anterior margin and constricted in
the anal cell, extending from base of third posterior
cell to tip of anal cell; preapical band and central band
broadly connected in whole of first posterior cell and
anterior part of second posterior cell, central band and
basal band connected in whole of discal cell; except for
hyaline base five hyaline spots, one from tip of costal
cell extending to fourth vein, one from tip of anal cell
to wing margin, one narrow one extending from ante-
rior margin to third vein at the level of posterior
crossvein, one basally in second posterior cell and one
occupying the apical tenth of the wing; wing almost
uniformly covered by microtrichia, only a tiny
glabrous section in base of costal cell.
Legs. - Front leg brown with blackish brown coxa,
tibia and metatarsus, paler other tarsi, pollinose ante-
riorly and basally on coxa 1 and on inner side of fe-
mur; mid leg brown with whitish basal two-fifth of
femur and dark apex of femur; hind leg brown with
basal eighth of femur whitish; femur 1 slender in 9
(ratio of length/ width 5.3, range 5.0-5.7) and slender
in 6 (ratio of length/width 5.3 ± se 0.1, range 5.2-
5.4); tubercles on distal three-quarters, inner row in
9 with 29.0 tubercles ± 0.8 (range 22-29) and in S
with 24.7 tubercles ±1.8 (range 19-29), outer row in
9 with 22.5 tubercles ± 0.9 (range 20-27) and in 6
with 22.5 tubercles ±1.7 (range 17-27).
Preabdomen. — Dorsally blackish brown, pollinose,
base more whitish pollinose; terga 2 and 3 anterolat-
erally with whitish pollinose spots; tip (centre of ter-
gum 4 and tergum 5) also whitish pollinose (fig. 64);
syntergum consisting of terga 1, 2 and 3, seam be-
tween terga 2 and 3 distinct; sternum 1 dark brown,
pollinose; sternum 1 basally fused to syntergum.
Female postabdomen. - Deflexed, terga 6, 7 and 8
rectangular plates (fig. 65); basal half of tergum 8 (fig.
66) glabrous; tergum 10 with four pairs of hairs, cer-
ei broad, ratio of length/width 1.9, covered with mi-
crotrichia and a number of hairs (fig. 66); sterna 5
and 6 single rectangular sclerites (fig. 65); sternum 7
angular anteriorly and constricted medially posterior-
ly; sternum 8 a single, triangular to V-shaped sclerite;
spiracle 7 in membrane; subanal plate (figs. 66-67)
somewhat pentagonal with rounded corners, posteri-
orly nine pairs of hairs; spermathecae (fig. 68) round-
ed and smooth; genital ring (fig. 69) rounded.
Male postabdomen. - Terga 5 and 6 single plates;
sterna 4 and 5 single rectangular sclerites; sternum 6
represented by two small triangular plates; sternum
7+8 a very short, band-like sclerite; spiracles 5 and 6
in membrane, spiracle 7 located rather dorsally, ante-
riorly of sternum 7+8; periandrium (fig. 70) rounded,
with about 20 pairs of hairs, covered with mi-
crotrichia; gonostyli rectangular in lateral view (fig.
9), ratio length/width 2.3, in posterior view apically
pointed, distal third on outer side covered with mi-
crotrichia, on inner side glabrous with only mi-
crotrichia on apical edge, on outer side some short
hairs on apical half; gonostyli interconnected via thin
processus longi; cerei very slender, ratio length/width
7.6, somewhat triangular, covered with microtrichia
and hairs; phallapodeme (fig. 71) rather slender, ante-
rior arm hardly broadening anteriorly and hardly
longer than posterior arm; ejaculatory apodeme (fig.
72) somewhat abruptly broadening anteriorly.
Diagnosis
Eurydiopsis subnotata gives its name to the subnota-
ta complex and can be recognized by its wingpattern
(almost hyaline apex, very small hyaline spot in mar-
ginal and submarginal cell), almost uniform distribu-
tion of microtrichia on the wing, depression in front
of ocellar tubercle, pollinose frons, minuscule rvB,
medium-sized ovb, small but distinct facial teeth,
slender front femora, long scutellar spines (25% of
body length), syntergum 1+2+3, rectangular 9 ter-
gum 8, medially constricted 9 sternum 7, pentagonal
subanal plate, broad female cerei, single male sternum
5, rectangular gonostyli with ratio length/width of
2.3, distal third of outer side of gonostyli covered with
microtrichia, slender and somewhat triangular cT cer-
ei and phallapodeme with almost equal-sized arms.
Acknowledgements
I'm grateful to the following curators for the op-
portunity offered to study Eurydiopsis in their collec-
tions: A. Albrecht (uzmh), B. Brugge (zma), N. E.
Evenhuis (bpbm), P. Grootaert (irsnb), P. J. van
Helsdingen (rmnh), G. C. McGavin (umo), B.
Pitkin (bmnh) and M. Dorn (mluh, Halle). D.
Burkhardt and I. de la Motte (University of Regens-
burg) placed material from Malaya and Sarawak at
my disposal. Comments on the manuscript by M.
Kotbra were highly appreciated.
239
Tijdschrift voor Entomologie, volume i4i, 1998
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Feijen, H. R., 1989. Diopsidae. - Cyclorrhapha III
(Schizophora other than Calyptratae). - In: G. C. D.
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Feijen, H. R., 1998. Teleopsis Rondani (Diptera, Diopsi-
dae): generic review and the ferruginea group from Sri
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Frey, R., 1928. Philippinische Dipteren. V. Farn. Diopsi-
dae. - Notulae Entomologicae 8: 69-77.
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bigas insectorum sistens, etc. 7 pp. Upsaliae.
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Osten Sacken, C. R., 1882. Diptera from the Philippine Is-
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mologische Zeitschrift 26: 83-120, 187-252.
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l'Institut Scientifique de Madagascar (A) 3: 65-76.
Shillito, J. F., 1971. The genera of Diopsidae (Insecta:
Diptera). — Zoological Journal of the Linnean Society 50:
287-295.
Steyskal, G., 1972. A catalogue of species and key to the
genera of the family Diopsidae (Diptera: Acalyptratae). -
Stuttgarter Beiträge zur Naturkunde, Serie A (Biologie),
No. 234. 20 pp.
Steyskal, G., 1975. Family Diopsidae. Pp. 32-36. - In: M.
Delfinado & D. E. Hardy (eds.). A catalog of the Diptera
of the Oriental Region. 3. Univ. Press Hawaii, Honolulu.
Tan, K. B., 1965. The taxonomy, biology, ecology and be-
haviour of some Malayan Diopsidae (Diptera). — M.Sc.
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(Diptera) or the 'Stalked-eye flies'. - The Malayan Scien-
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Diopsidae nouveau de Madagascar. Revue de Zoologie et
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40: 181-198, plate 1.
Received: 18 August 1998
Accepted: 12 November 1998
240
L. MATILE
Muséum national d'Histoire naturelle, Laboratoire d'Entomologie et EP 90 du CNRS
FIRST PALAEARCTIC RECORD OF THE GENUS
LAURYPTA EDWARDS (DIPTERA: KEROPLATIDAE)
Matile, L., 1999. First Palaearctic record of the genus Laurypta Edwards (Diptera: Keroplati-
dae). - Tijdschrift voor Entomologie 141 [1998]: 241-244, figs. 1-5.[issn 0040-7496]. Pub-
lished 1 March 1999.
The genus Laurypta Edwards is recorded for the first time from the Palaearctic region, where it
is represented by rwo species, L. exserta Ostroverchova (comb, n.), from the Russian Far East,
and L. tripotini sp. n. from South Korea, which is described and illustrated. Laurypta belongs to
the afrotropical-oriental track, which has extensions to the Eastern Palaearctic and the North-
ern Australasian. No other genus of Keroplatidae follows this same track, which can be dated
from the Miocene.
Correspondence: Loïc Matile, Laboratoire d'Entomologie, Muséum national d'Histoire na-
turelle, F-75005 Paris, France. E-mail: lmatile@mnhn.fr.
Key words. - Diptera; Keroplatidae; Palaearctic; new species; new combination; biogeogra-
phy; afrotropical-oriental track.
Laurypta was described by Edwards ( 1 929) as a sub-
genus of ' Platyura Meigen', now Orfelia Costa. The
species included in the subgenus were Platyura leptura
Edwards, the type species, from the Malay Peninsula,
P. tripunctata Senior- White, from Sri Lanka, and P.
laevis Enderlein, from the Seychelles. Colless (1966)
described Orfelia (Laurypta) apicalis from Palau Is-
and, thus recording Laurypta for the first time from
the Australasian Region. Matile (1970) added a fur-
ther Afrotropical species, Orfelia (Laurypta) scalaris
from Cameroun, and later recorded it from Central
African Republic (Matile 1974). Laurypta was, formal-
ly erected at generic rank by Matile (1988a) in a paper
recording L. scalaris from Ivory Coast and Congo, and
citing an undescribed species from Senegal. The genus
had thus up to now an Afro-oriental distribution with
an extension in the Australasian. In a collection of
Keroplatidae from south Korea recently sent by M.
Pierre Tripotin, stood several specimens of Orfeliini
belonging to an undescribed species of this genus.
This is the first record of Laurypta for the Palaearctic
Region. The distribution of the genus is given in fig. 1 .
As already pointed out (Matile 1988a), Laurypta is
well characterized by a number of apomorphic traits
such as the loss of the veins Sc, and An and of the out-
er tibial spurs, the regular disposition of the tibial setu-
lae, the reduction in number of the tibial setae, the
mediotergite bearing a group of strong bristles, but
none of these characters are exclusive for the genus in-
side the tribe Orfeliini. On the other hand, the leng-
thening of the male tergite DC, cerei and hypoproct,
and also of the gonostyles, these four parts each being
longer than the gonocoxal synsclerite, may be consid-
ered autapomorphic for the genus (for a cladistic analy-
sis of the characters of the Keroplatidae, see Matile
1990). Pending a generic revision of the Orfeliini, the
sister-group of Laurypta cannot be ascertained.
The new species described here is very close to
Platyura exserta Ostroverchova, 1979, from the Russ-
ian Far East (Sikhote-Alin Reserve, Russian Maritime
Province), which was later attributed to Pyratula Ed-
wards (Krivosheina & Mamaev, 1988), and keyed in
this genus by Zaitzev (1994). This species also be-
longs to Laurypta [comb. n. : Platyura (Z.) exserta Os-
troverchova, 1979: 40 and plate 8, fig. 5].
Description
Laurypta tripotini sp. n.
(figs. 2-5)
Type material. - Male holotype: South Korea,
Posok-sa temple (Keumsan), Malaise trap, 30.05.-
4.06.1998. -Paratypes: 2 ó\ 39; \6 and 2$: same
data as holotype. 1 6: same data but 26-30.05.1998.
1 9 : Sanan Hi (Keumsan), very dry clearing, Malaise
trap, 20.viii.1997. All material, collected by P. Tripo-
tin, in Muséum national d'Histoire naturelle, Paris.
Male holotype. - Length of wing: 2.9 mm. Head:
occiput ochreous with a silvery pruinosity according
to angle of light. Three ocelli, the outer ones far from
eye margin, ocellar callus black brown. Antennae:
scape, pedicel and basis of first flagellomere yellow,
241
Tijdschrift voor Entomologie, volume mi, 1998
scalaris
Fig. 1 . - Distribution of the genus Laurypta.
rest of flagellum black brown. Face and clypeus
ochreous, mouthparts and palpi black brown.
Thorax: humeral calli pale yellow. Scutum ochreous,
bearing four small, indistinct, prescutellar brownish
spots. Scutellum brown, mediotergite yellow, brown in
middle at level of insertion of macrochaetae. Lateroter-
gite brown dorsally, pleura yellow. A small brownish
spot on the mesanepisternite, at level of insertion of a
group of minute hairs. Metepisternite also with a patch
of short hairs.
Legs yellow, tibial spurs 1:1:1, black. Tibia and
protarsus I of equal length.
Wings yellow, with a faint apical spot under apex
of R,. C reaching the tip of the wing, strongly pro-
duced after R,. Sc very short, ending in costa well be-
fore Rs, Sc, absent. R4 very short, ending after the
middle of the R,-R, interval (3:2.4). Radiomedian fu-
sion a little shorter than Rs. Anal absent. Haltères yel-
low, darkened at tip.
Abdomen: tergite I ochreous, the following tergites
ochreous with a wide apical brown band. Sternites
ochreous. Terminalia: tergite IX and synsclerite yel-
low, proctiger and gonostyles brown.
Male genitalia (figs. 2-4). - Ninth tergite long,
wider in the basal third, extending far distad of the
gonocoxite margin. Cerci long and narrow, extending
distad of apex of gonostyles, hypoproct shorter, not
reaching apex of cerei. Gonocoxites entirely fused ven-
trally, the ventral margin of the synsclerite smoothly
curved inwards. Synsclerite entirely open dorsally, the
internal margin bearing, on each side, a short process
bearing four closely appressed short spines. Gonos-
tyles bifid, the ventral lobe wide, the dorsal one nar-
row and bearing at the internal margin rows of
stronger and shorter setae. Aedeagus small and weakly
sclerotized.
Female as the male, but the brown apical abdomi-
nal bands narrower. Terminalia brown black, cerei
small, one-segmented, blunt at apex.
Remarks. - The male genitalia are of the usual Lau-
rypta type (see drawings by Edwards 1928, for L. lep-
tura, and Matile, 1988a for L. scalaris), and obviously
very close to those of L. exserta (Ostr.). The types of
the species described by Ostroverchova cannot be bor-
rowed (Ostroverchova pers. comm.), but according to
the sketch she gave of this species, L. tripotini differs
by the ninth tergite wider at base (compare fig. 4, L.
tripotini drawn under the same angle and magnifica-
tion than fig. 5, L. exserta after Ostroverchova), and
the gonostyles wider and straighten The Korean
species further differs from L. exserta by the scutum
ochreous instead of brown black and the abdominal
tergites V-VII banded instead of uniformly dark. Os-
troverchova also mentions the presence of Sc, and of
an anal vein 'well developed but not reaching the wing
margin' - as stated above, the absence of Sc2 and An
are diagnostic of the genus Laurypta, and the two
species are so close in male genital structure that I sus-
pect a confusion between Sc2 and humeral crossvein h,
and between An and Cu2 (on the other hand, these ve-
nation characters would check with the previous as-
signment of exserta in Pyratula).
242
Matile: First Palaearctic Laurypta
Figs. 2-5. Laurypta species —
2, L. tripotini sp. n., male
holotype, postabdomen, lat-
eral; 3, ditto, dorsal; 4, ditto,
proctiger and ninth tergite,
dorsal; 5, L. exserta (Ostr.),
proctiger and ninth tergite,
dorsal (after Ostroverchova).
0.3 mm
Discussion
In his diagnosis of Laurypta, Edwards ( 1 929) states
the mesanepisternite as bare. L. laevis, scalaris, leptura,
apicalis and the undescribed species from Senegal all
show the patch of minute mesanespisternal hairs men-
tioned in the description of L. tripotini. The presence
of metepisternal hairs has not been noted either by Ed-
wards and the diagnosis of the genus should thus be
amended.
In the keys to the genera of Keroplatinae given by
Hutson et al. (1980) or Zaitzev (1994), Laurypta
would come close to the genus Orfelia Costa, from
which it differs conspicuously by the absence of outer
tibial spurs, by the rows of tibial setulae all alike and
the anal vein absent.
L. apicalis (Colless) , described on a single damaged
female, differs strikingly from the other Laurypta by
its pictured wings, the pattern of which resembles the
New Caledonian Dimorphelia Matile. The holotype
was kindly examined for me by Dr. Raymond J. Gag-
né (usnm): there are 14 flagellomeres (normally 12 in
females of Dimorphelia, but I have seen specimens
with 13), and there are anepisternal setae which es-
caped Colless attention. The mouthparts are pointed,
but shorter than the combined length of the first two
palpomeres (reaching the last palpomere in Dimor-
phelia), the anal vein is absent (faint in Dimorphelia)
and the metepisternite is bare, so that L. apicalis is
most probably correctly placed in the genus.
The genus Laurypta thus follows an afro-oriental
track, with two extensions, one Eastern Palaearctic, the
other Australasian. The presence of a few Oriental ele-
ments in the Japanese and Eastern Asian fungus-gnat
fauna has been signalled by Okada (1938a, b,1939),
while discussing certain species. The relations between
afrotropical, oriental and australasian Sciaroidea have
been briefly discussed by Matile (such as 1988a, b,
1990 and 1992). On more general grounds Croizat
(1958, 1964) has stressed the importance of an 'axis of
dispersal: 'Eastern Siberia/Japan-Java, etc. [with ...] an
East African outlier' (1964: 512), which is typically
that of Laurypta, with lacunae which will probably be
243
Tijdschrift voor Entomologie, volume mi, 1998
filled in the future.
The Oriental-australasian track corresponds to two
periods, Miocene and Pliocene-Pleistocene (Holloway
& Jardine 1968); the fact that the distribution of Lau-
rypta includes also the Afrotropical region implies that
the genus is at least of Miocene origin; the extension
to Eastern Asia could be as recent as the Pliocene.
Of the 49 known genera of Orfeliini, 27 are en-
demic to one biogeographical region, 4 are Holarctic
[Asindulum Latreille, Macrorrhyncha Winnertz, Pa-
laeoplatyura Meunier (living in North America, fossil
in Baltic amber) and Platyura Meigen], 4 are Austro-
neotropical {PLtnarivora Hickman, Pyrtaula Ed-
wards, Rypatula Edwards and Taulyrpa Edwards), one
Afro-oriental {Ralytupa Edwards) and one Afro-neo-
tropical {Lyprauta Edwards, but there may be a Palae-
arctic species, Platyura oberthueri Matile). Pyrtulina
Matile is Afro-australasian, with probable Oriental
representatives. The other 1 1 genera cover at least
three biogeographical regions, but Laurypta is the
only one showing the distribution discussed above,
although a phylogenetic analysis of certain genera
may in the future reveal such relationships among
species-groups inside genera. None of the genera of
Macrocerinae or of Keroplatinae Keroplatini show a
Laurypta-like distribution. The 'oriental group' of the
genus Heteropterna does follow a similar track, but its
phylogenetic relationships are unsatisfactorily re-
solved (Matile 1990). Although there are a few other
Oriental elements in the far Eastern Palaearctic Scia-
roidea (Okada op. cit., Matile unpubl. data), these re-
main therefore anecdotal.
Acknowledgments
It is a pleasure to acknowledge the generosity of M.
Pierre Tripotin in giving his very interesting material
of Korean Diptera and other orders to the Muséum
national d'Histoire naturelle, Paris. I am very grateful
to my friends Dr Pavel Putshkov (Institute of Zoolo-
gy, Kiev), for translating into French the original de-
scription of Laurypta exserta Ostr. during his last stay
in Paris, and Dr Raymond J. Gagné (usnm) for notes
taken on the holotype of Orfelia apicalis Colless.
References
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Received: 15 August 1998
Accepted: 12 November 1998
244
Volume 141
1998
/
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Tijdschrift
voor
Entomologie
A journal of systematic and evolutionary
entomology since 1858
f*.
^f-
<?f^\
Netherlands Journal of Entomology
Published by the Netherlands Entomological Society
Tijdschrift voor Entomologie
A journal of systematic and evolutionary entomology since 1858
Scope
The Tijdschrift voor Entomologie' (Netherlands Journal of Entomology) has a long
tradition in the publication of original papers on insect taxonomy and systematics.
The editors particularly invite papers on the insect fauna of the Palaearctic and
Indo-Australian regions, especially those including evolutionary aspects e.g.
phylogeny and biogeography, or ethology and ecology as far as meaningful for
insect taxonomy. Authors wishing to submit papers on disciplines related to
taxonomy, e.g. descriptive aspects of morphology, ethology, ecology and applied
entomology, are requested to contact the editorial board before submitting.
Usually, such papers will only be published when space allows.
Editors
E. J. van Nieukerken (elected 1986) and M. Schilthuizen (1998)
Co-editors
A. W. M. Mol (1990) and R. T. A. Schouten (1990) and P. L Th. Beuk (1998)
Advisory board
M. Brancucci (Basel), N. E. Stork (London) and M. R. Wilson (Cardiff).
The 'Tijdschrift voor Entomologie' is published in two issues annually by the
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Published with index of volume 141 (1998).
Graphic design
Ontwerpers B.V., Aad Derwort, 's-Gravenhage
Tijdschrift voor Entomologie
Contents of volume 141
Articles
129 Anderson, R.S.
New species of Sicoderus Vanin from the Virgin Islands (Coleoptera: Curculionidae;
Curculioninae; Otidocephalini).
I Bickel, DJ.
Australian, Melanesian, and Micronesian Acropsilus Mik (Diptera: Dolichopodidae).
137 Chen, P.P. & H. Zettel
A taxonomie revision of the oriental water strider genus Ventidius Distant
(Hemiptera, Gerromorpha, Gerridae).
19 Choi, S.-W.
Systematics of the genus Heterothera Inoue (Lepidoptera, Geometridae: Larentiinae)
209 Epler, j.H. , A.D. Harrison & L. Hare
Acinoretracus, a new Afrotropical genus for some species previously placed in
Dicrotendipes (Diptera: Chironomidae: Chironominae).
49 Feijen, H.R.
Teleopsis Rondani (Diptera, Diopsidae): generic review and the ferruginea group from
Sri Lanka.
221 Feijen, H.R.
A revision of Eurydiopsis Frey (Diptera, Diopsidae) with description of four new
Oriental species.
Hare: see Epler
Harrison: see Epler
65 Ingrisch, S.
A review of the Elimaeini of Western Indonesia, Malay Peninsula and Thailand (Ensife-
ra, Tettigoniidae, Phaneropterinae).
Karsholt: see Rutten
241 Matite, L.
First Palaearctic record of the genus Laurypta Edwards (Diptera: Keroplatidae).
109 Rutten, T. & O. Karsholt
Bryotropha mundella (Douglas): a new synonym of Bryotropha umbrosella (Zeiler)
(Lepidoptera Gelechiidae).
115 Oswald, J.D.
Annotated catalogue of the Dilaridae (Insecta: Neuroptera) of the world.
Zettel: see Chen
Tijdschrift voor Entomologie, volume i4i, 1998
Book reviews
18 Zlata S. Gershenson & Sandrine A. Ulenberg, 1998. The Yponomeutinae (Lepido-
ptera) of the World exclusive of the Americas. • Menno Schilthuizen & Henk Vallen-
duuk, 1998. Kevers op kadavers. «Johan van Zoest (ed.), 1998. Biodiversiteit. [E. J.
van Nieukerken]
48 Orthoptera sounds: D.R. Ragge & W.J. Reynolds, 1998. The songs of the Grasshoppers
and Crickets of Western Europe. • D.R. Ragge & W.J. Reynolds, 1998. A sound
guide to the Grasshoppers and Crickets of Western Europe. • H. Bellmann, 1993a.
Heuschrecken beobachten, bestimmen. • H. Bellmann, 1993b. Die Stimmen der
heimischen Heuschrecken. • H. Bellmann & G. Luquet, 1995. Guide des sauterelles,
grillons et criquets d'Europe occidentale. • F.-R. Bonnet, 1995. Guide sonore des
sauterelles, grillons et criquets d'Europe occidentale. • R.M.J.C. Kleukers, E.J. van
Nieukerken, B. Odé, L.P.M. Willemse & W.K.R.E. van Wingerden, 1997. De
sprinkhanen en krekels van Nederland (Orthoptera). • B. Odé, 1997. De zingende
sprinkhanen en krekels van de Benelux. [E. J. van Nieukerken]
1 36 F. Bos & M. Wasscher, 1997. Veldgids libellen. - Stichting Uitgeverij knnv, Utrecht
[J. van Tol]
Reviewers for volume 1 4 1
U. Aspöck (Wien, Austria), P. J. Chandler (Slough, UK), K. D. Dijkstra (Leiden), P. Grootaert
(Brussel, Belgium), J. Holloway (London, UK), R. de Jong (Leiden), M. Kotbra (Berlin,
Germany), R. Oberprieler (Canberra, Australia), D. A. Polhemus (Honolulu, USA), D. Rentz
(Canberra, Australia), O. A. Saether (Bergen, Norway), Z.-Q. Zhang (London, UK).
Dates of Publication
Volume 141 (I), pages 1-128, 30 November 1998
Volume 141 (2), pages 1 29-244, i-viii, I March 1999
Nederlandse Entomologische Vereniging, Amsterdam ISSN 0040-7496
NEW TAXA DESCRIBED IN
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 141
COLEOPTERA
Sicoderus hirsutiventris Anderson, 1999 131
Sicoderus ivieorum Anderson, 1999 129
Sicoderus panini Anderson, 1 999 133
Diptera
Acinoretracus Epler, Harrison & Hare, 1999 ... 210
Acropsilus albitibia Bickel, 1998 13
Acropsilus boharti Bickel, 1998 12
Acropsilus colmani Bickel, 1998 8
Acropsilus kuranda Bickel, 1998 8
Acropsilus malaita Bickel, 1998 10
Acropsilus maprik Bickel, 1998 12
Acropsilus nigricornis Bickel, 1998 13
Acropsilus putosa Bickel, 1998 16
Acropsilus toma Bickel, 1998 10
Acropsilus Wo? Bickel, 1998 16
Eurydiopsis brevispinus Feijen, 1999 229
Eurydiopsis glabrostylus F eijen, 1999 232
Eurydiopsis helsdingeni Feijen, 1999 233
Eurydiopsis sarawakensis Feijen, 1999 236
Laurypta tripotini Matile, 1999 241
Teleopsis krombeini Feijen, 1998 57
Teleopsis maculata Feijen, 1998 61
Heteroptera
Ventidius(s. str.) longitarsus Chen & Zettel, 1999. .
155
Ventidius(s. str.) pilosus Chen & Zettel, 1999. . 169
Ventidius (s. str.) polhemorum Chen & Zettel, 1999 .
165
Ventidius (Ventidioides) beissi Chen & Zettel, 1999.
197
Ventidius {Ventidioides) nieseri Chen & Zettel, 1999
196
Ventidius (Ventidiopsis) yangae Chen & Zettel, 1999
203
Lepidoptera
Heterothera distinctata Choi, 1998 33
Heterothera eclinosis Choi, 1998 35
Heterothera hoenei Choi, 1998 28
Heterothera kurenzovi Choi, Viidalepp & Vasjurin,
1998 42
Heterothera mussooriensis Choi, 1998 27
Heterothera obscurata Choi, 1998 31
Heterothera stamineata Choi, 1998 36
Heterothera triangulata Choi, 1998 36
Heterothera yunnanensis Choi, 1998 33
Orthoptera
Elimaea (Elimaea) nautica Ingrisch, 1998 87
Elimaea (Elimaea) ?/wz'z' Ingrisch, 1998 86
Elimaea (Rhaebelimaea) apicata Ingrisch, 1998. . . 81
Elimaea (Rhaebelimaea) maninjauensis Ingrisch,
1998 79
Elimaea (Rhaebelimaea) mentaweii Ingrisch, 1998 79
Elimaea (Rhaebelimaea) modiglianii Ingrisch, 1998 .
80
Elimaea (Rhaebelimaea) pentaspina Ingrisch, 1998 . .
82
Elimaea (Rhaebelimaea) pseudochloris Ingrisch, 1998
81
Elimaea (Rhaebelimaea) sinuata Ingrisch, 1998. . . 82
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Tijdschrift voor Entomologie
Volume 141, no. 2 [1998]
Articles
129 R.S.Anderson
New species of Sìcoderus Vanin from the Virgin Islands (Coleoptera:
Curculionidae; Curculioninae; Otidocephalini).
137 P.P. Chen & H. Zettel
A taxonomie revision of the oriental water strider genus Ventidius Distant
(Hemiptera, Gerromorpha, Gerridae).
209 J.H. Epler, A.D. Harrison & L. Hare
Acinoretracus, a new Afrotropical genus for some species previously
placed in Dicrotendipes (Diptera: Chironomidae: Chironominae).
221 H.R. Feijen
A revision of Eurydiopsis Frey (Diptera, Diopsidae) with description of
four new Oriental species.
241 L. Matile
First Palaearctic record of the genus Laurypta Edwards (Diptera:
Keroplatidae).
Book reviews
I 36 F. Bos & M. Wasscher, 1 997. Veldgids libellen. - Stichting Uitgeverij KNNV, Utrecht.
[). van Tol]
© Nederlandse Entomologische Vereniging, Amsterdam
Published I March 1999 ISSN 0040-7496
ERNST MAYR LIBRARY
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